Last Alphas 02.

0 - Claimed by the Hill

Wolf Marina Maddix
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CLAIMED BY THE HILL WOLF

THE LAST ALPHAS BOOK 2

MARINA MADDIX
 CONTENTS

About This Book

Chapter 1
Chapter 2
Chapter 3
Chapter 4
Chapter 5
Chapter 6
Chapter 7
Chapter 8
Chapter 9
Chapter 10
Chapter 11
Chapter 12
Chapter 13
Chapter 14
Chapter 15
Chapter 16
Chapter 17
Chapter 18
Chapter 19
Chapter 20
Chapter 21
Chapter 22
Chapter 23
Chapter 24
Chapter 25
Chapter 26
Chapter 27
Chapter 28
Chapter 29
Excerpt from Fated to the Rogue Wolf
Also by Marina Maddix
About the Author
 ABOUT THIS BOOK

She just wants to do her job. He just wants her.

Since being taken in by the wolf shifters of the Valley, my sole focus has
been curing their fertility problem. I don’t care that everyone seems to think I
should accept the bite that would transform me into one of them. I have more
important things to think about.
Then I meet him — the dead-sexy new alpha of the Hill wolves who can’t
seem to get enough of my curves. Markon stirs something in me that I’ve
never felt before, and that kind of distraction is the last thing I need.
Unfortunately, it’s all I can think about.
And now he’s pressuring me to become a wolf, too. Something about finding
out if I’m his fated mate. Nonsense! Fated mates don’t actually exist.
Do they?

Claimed by the Hill Wolf is Book 2 in The Last Alphas trilogy. Read an
excerpt of Fated to the Rogue Wolf at the end of this book.

THE LAST ALPHAS

Mated to the Forest Wolf
Claimed by the Hill Wolf
 Fated to the Rogue Wolf

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more!
 1
NATALIE

“W ho wants to hear the story of how Aunty Sienna, Aunty Natalie

and I came here?”
Every tiny hand within the sound of Arlynn’s voice shot up. Sienna’s dark
curls bobbed when she looked up from the wobble ring game she was playing
with a group. She loved telling the story, and why shouldn’t she? It turned
out pretty well for her.
The children at my finger-painting table jumped up and ran to Arlynn,
abandoning the paints I’d painstakingly mixed up using berries, plants and
bugs found in the forest. I couldn’t blame them, really. Kids didn’t respond
well to my habit of being logical, but they loved her free-spirited ways.
That was partly why she suggested this children's’ fair in the first place. The
kids had all loved her the minute we stepped foot in this village, even though
we were all Terran. And when Sienna became the mate of the tribe’s alpha,
the kids adored her, too.
Me, not so much. At best they tolerated my presence; at worst, they looked at
me like I was an alien. Which I guess was pretty accurate, from their point of
view.
Even though my tall, curvy frame more closely resembled female Wargs, my
long, blonde hair and blue eyes were oddities among the tribe of wolf-like
shifters. Of course, they accepted Arlynn’s red and purple-dyed hair without
question, even though her dark brown roots were finally growing out.
The three of us sat side by side on a log as the children gathered around us,
their glowing faces gazing up at us expectantly. They all knew the story, but
they couldn’t wait to hear it again.
“There we were, being loaded on a Terran shuttle like livestock,” Arlynn
started in a spooky voice. Immediately, little Blen raised his hand.
“What’s livestock?”
“Oh, that’s a word for the animals that we raise. Like cabras.”
The little ones ‘aahed’ and Arlynn continued.
“The people at the Training Center told us we were going on exciting, secret
missions, but they lied.”
Big amber, brown, and forest-green eyes grew wide. “How did you find out?”
asked Eileah, one of the few girls to be born in recent years. Sienna had saved
her from choking on our first night in the village, an act that turned Warg
opinion of us ‘alien interlopers’ from fear to gratitude.
Sienna leaned forward. “I overheard the guards talking. They said we were to
become slaves!”
Gasps all around. Of course, it would have been inappropriate to tell them the
whole truth — that we’d been destined to become sex slaves to military
camps.
“So when our guards least expected it, we overpowered them and Natalie
landed the shuttle in Hill Warg territory.”
All eyes turned to me, waiting for me to take over.
“You flew a sky bird?” asked another little boy, Frink. “What was it like?”
I glanced at Arlynn and Sienna. They both nodded and smiled, as if
interacting with children was the easiest thing in the world.
“Well, um, it’s, um…it was like…flying.”
They blinked at me, unimpressed.
“Did you know how?” Frink pressed.
“No, but I’d studied enough aerophysics and mechanical engineering at the
Center to understand the basic principles. It was a simple matter of
extrapolating the dynamics between the instrumentation of the aircraft and
the physical properties of its exterior, then calculating the lift produced by the
sub-boosters.”
Crickets.
Way to go, Nat. You bored little kids into muteness.
“What she means to say,” Arlynn said, “is that she didn’t know how to fly it
at all but she did it anyway. It was scary!”
The kids ‘oohed’ this time.
“Then what happened?”
“Well, then we ran from the sky bird — we call it a ‘shuttle’ — and right into
the clutches of a great, big grumpus!”
More gasps, and this time they actually looked frightened. Totally irrational.
Since we were sitting in front of them telling this tale, basic logic dictated
that everything turned out fine.
“That’s when the brave, strong alpha of the Valley Warg saved us!”
“Yay!” they all shouted.
Sienna picked up the story. “But I didn’t know Solan was a good Warg at the
time, so I grabbed a stick and bonked him on the nose.”
I had to admit that little kid laughter was pretty contagious.
“Of course, you all remember our early days in the village, right? Thank
goodness Wargs can boost their speech so aliens like us can understand them.
Otherwise, Blen, you might not have been able to teach me how to play
Catch-A-Mate with the wobble rings.”
Blen nodded vigorously. “And Solan caught your ring! That meant he was
your fated mate.”
I couldn’t help snorting. Sienna shot me a dirty look and continued.
“That’s right. But we didn’t know that at the time, either.”
“Is that when the mean Hill tribe kidnapped you?” Frink asked.
Sienna paused. Solan was working hard to unite the Valley and Hill tribes.
They’d split generations earlier and had been feuding ever since. The task of
reversing generations of hatred was proving difficult, even with the wee ones.
She chose her words carefully.
“It was all a misunderstanding,” she finally said. “But yes, that’s when we
were taken to the Hill tribe’s village.”
The truth would curdle their blood. Even someone as clueless as I was
regarding children knew the whole story would give them nightmares for
weeks.
“And that’s where Solan gave me the bite that turned me into one of you!”
They all cheered.
“And I was also given the bite that changed me,” Arlynn interjected. She
conveniently left out that the bite had been administered by the Hill tribe’s
alpha against her will.
More cheering. Then they all looked at me.
“Why aren’t you a Warg, Aunty Nat?” Eileah asked.
I cleared my throat. “Well, I’m currently working with Jorek on a solution to
the tribe’s unusual fertility problem. We’re making progress and I don’t want
to risk breaking the streak. You see, as far as I can tell, when certain chemical
compounds in a Warg’s saliva mix with Terran blood, a metamorphosis of
the putative central neurotransmitters in the medulla oblongata occurs—“
“What she means,” Arlynn interrupted, “is that she’s trying to find a way for
you all to have baby sisters very soon. She’s worried that the change might
make her brain think differently.”
“But don’t you want to find your fated mate?” Eileah asked, her tone
strangely worried.
I sighed, rolling my eyes. “I don’t believe in—“
Sienna jabbed an elbow in my side. The dark look she shot me practically
growled, ‘Don’t you dare.’
“Let’s just say that finding a mate—“
“Your fated mate,” Blen corrected.
“Whatever. It’s not important for me.”
Eileah’s eyes grew wide. “Don’t you want a mate?”
“Of course she does,” Arlynn said quickly, smiling. “She just doesn’t know it
yet.”
The children grinned up at me, like they were in on a secret joke. As much as
I liked their change in attitude toward me, I didn’t approve of lying to kids.
I’d been lied to my entire life, and I refused to continue the cycle.
“No, really. I’m far too busy to worry about men and mates and all of that
nonsense. Once I solve the mystery of why female offspring are so rare, I’ll
accept the bite and become a Warg. If I fall in love, fine. If not, no big deal.”
Sienna and Arlynn shared a glance that spoke volumes. As if they knew
better than me what I wanted from my life. Ridiculous! I’d known that I
wanted to be a respected scientist long before my pathetic excuses for parents
abandoned me the Training Center. Love never entered into the plans for my
future, and that went double for so-called ‘fated mates’.
“Do we believe her?” Arlynn asked the kids.
“NO!” They all doubled over in giggles, pointing tiny fingers at me.
“It’s true!” I objected, but no one could hear me over the laughter. “Really!”
Why wouldn’t they believe me? Arlynn might have been desperate to find a
mate, but not me. I didn’t care one way or the other.
Really.
 2
MARKON

“M arkon, how can our tribe follow Thrane after his humiliation in
battle?”
Thrane tensed next to me, and I didn’t need to look at him to know he was
grinding his teeth. It sounded like he was chewing on bonknut shells.
The tribal council stared down at us from their perch on the platform in the
village’s meeting hall. Normally, my brother enjoyed that honored spot
because it enhanced his position of power as the Hill tribe’s alpha. As his
second in command, I always stood one step behind him.
Now we were the ones being judged. Thankfully, it was a closed session.
“Wise council of the Hill Warg,” I said, keeping my voice calm and
agreeable, “it cannot be denied that Solan breeched etiquette by allowing
Thrane to live after defeating him in the Holmgang challenge he called. As
the alpha of the Valley Warg, Solan knew better than anyone the
consequences of such an action.”
“Bastard!” Thrane barked. I shot him a warning glance.
“Since the Great Split, it has been one of the few laws that both tribes agreed
on,” said Nabor, the oldest of the bunch. “If you respect the warrior you have
defeated in Holmgang, you honor him by taking his life.”
“Yes, I know, but—“
Nabor talked over me. “That means even Solan, a vicious Valley Warg,
couldn’t see any worth in Thrane. Why should we?”
“Don’t you fools see?” Thrane snapped. “This is exactly what he wants. By
disgracing me, he hopes to sow the seeds of discord in the Hill tribe. With me
out of the way, the Valley scum can wipe us out. You’re idiots if you can’t
see that!”
The council mumbled to each other, whispering and casting wary glances at
Thrane. He wasn’t helping his cause and, if I didn’t do something, they might
vote to cast him out, leaving me in a position I never expected or wanted.
Though it was the right of the tribal council to oust an alpha, it had never
been done in remembered time.
“Know what I think?” Thrane prowled back and forth in front of the
platform, glaring up at the council. He dragged a grubby hand through his
long, greasy black hair, giving me a view of the raw scar that ran around his
ear — the ear Solan had almost ripped clean off my brother’s head. I’d told
him to wash up before the meeting, but he ignored me, as usual.
After the battle with Solan’s tribe, Thrane wasn’t the same. He stopped taking
care of himself, letting his normally perfect appearance become bedraggled,
at best. He patrolled the perimeter of the village at all hours, only sleeping a
few hours a night and rarely eating. He certainly didn’t govern, leading more
and more tribe members to seek my counsel.
It took a few weeks, but the council eventually caught wind of it and called
this session to give him a chance to defend himself. Now I had the alarming
suspicion that Thrane was about to blow it to the next world and back.
“I think someone on the council has been leaking information to the Valley
scum.”
Shocked gasps echoed around the hall.
Oh shit!
I grabbed his arm to silence him, but he yanked free and continued glaring at
the men who would determine his fate. Our fates.
“What are you saying, Thrane?” Nabor demanded, his pale amber eyes
growing fiery.
“I think I was quite clear, Nabor. But if the words I used were too big for
your tiny brain to understand, let me put it another way. One of you is a spy.”
Roars of protest and fury rang through the building. One of the younger
council members, Pimmit, even tried to lunge at Thrane. The others
restrained him while I shoved Thrane backward and got in his face.
“What do you think you’re doing? You can’t accuse the council of spying.”
“Well, someone tipped off the Valley scum that we had his mate,” he
seethed. “Besides, I can say whatever I want. I’m the alpha of this tribe, in
case you’ve forgotten. Everyone else seems to have.”
Stubborn fool! This is why Thrane needed me as his second — to smooth
things over when he inevitably insulted someone.
“Just stand there and keep your mouth shut, do you hear me? I’m not going to
let you sabotage yourself and leave me stuck as the acting alpha.”
He glared at me but pressed his lips tight. Good boy.
I loved and respected Thrane, but it never failed to amaze me that we were
brothers. Aside from the large, muscled physiques we inherited from our
father, we looked nothing alike. Thrane was very proud of his thick, black
hair and flaming orange eyes. He often ribbed me about my short, brown hair
and eyes the color of sunlit leaves, claiming such a ‘pretty’ man could never
make an effective alpha. As always, I would smile and play the peaceable
diplomat to his volatile warrior.
Just as I was doing now.
Spinning around, purposely blocking the council’s view of my hotheaded
brother, I held my hands up to get their attention.
“Or,” I shouted, waiting for them to quiet down. One by one, they all turned
to me, waiting. I had a suspicion that they were looking for any excuse to
keep him or they would have exiled him by now. But he certainly wasn’t
making it easy.
“Or,” I continued, “Solan was right about the alien interlopers wanting to
destroy us.”
I let that sink in for a moment. I could practically hear hearts starting to race.
“If he is, then it would be in our best interests to agree to his suggestion to
unite our two tribes.”
The council sat stunned. Pimmet’s jaw dangled open as if he was trying to
capture a bug, and Simwat coughed furiously. Nabor was the first to find his
words.
“Markon, are you really suggesting that we unite with the Valley scum, our
mortal enemies since the Great Split?”
I rolled back my shoulders and steeled myself. “I am.”
Shouts and growls erupted again, but quieter this time. They’d never heard
such a preposterous suggestion, one voice said. I was a traitor to ask this of
them, said another. It was a ploy by the Valley scum, they just knew it, added
a third. I let them rant until it died down to a murmur.
“Council of the Hill Warg, you have served our tribe honorably and with
great mindfulness. Solan’s tribe has been our enemy for so long that nobody
alive remembers why.”
“It started back when our ancestors, Tooibas and Vanter, shared the role of
alpha, but then fought over which of them would take the most beautiful
female as his mate.“
“Yes, we all know the folklore, Simwat,” I interrupted. “But why are we
Solan’s enemy? Have you ever asked yourself that question? Of course not.
It’s always been that way and it always will be, right?”
A couple of the more dull-witted council members nodded eagerly. The
others remained mute, watching me thoughtfully. They were willing to listen,
which was more than I could say for Thrane. He snorted behind me. I could
almost see him rolling his eyes.
“So, what if he was right? What if the Terrans have plans to attack us, as he
claims? Remember, he has the three Terran females who might have inside
information.”
Thrane huffed again. “We had two of them.”
I barreled over him. “If what he says is true, we can’t afford to have two
enemies. We just don’t have the warriors to defend ourselves. But if we join
forces…”
“The Terrans haven’t tried to attack us for two generations,” Pimmit argued.
“Why would they do so now?”
I shrugged. “Maybe they want to expand their territory. Maybe they want to
rape the resources of the forest. Maybe they simply want to eradicate us once
and for all. The reasons don’t matter. What matters is that the threat is real.
We must take action before it’s too late.”
“You believe Solan’s claim, Markon?” Nabor asked.
I was suddenly and keenly aware that all eyes were on me. This wasn’t my
place. My place was supporting my brother, and that’s what I’d intended to
do. But now they were actually listening to me. They wanted to know my
opinion. Me!
Guilt washed over me that I’d somehow stolen Thrane’s spotlight, but a
small, glowing part that lived deep down inside felt even more guilty for
liking it.
“I do.”
Thrane shoved me aside. “I can’t believe what I’m hearing! Are you all
actually considering this?”
Nabor growled and let his beast come forth just enough to show Thrane that,
even at his advanced age, he was no one to be trifled with. Thrane stood his
ground but didn’t advance any farther.
“Your brother makes a good argument, Thrane. Only a fool would dismiss it
out of hand.”
Thrane threw his hands in the air. “Then I’m a fool. And you fools can all die
at the claws of the Valley scum, for all I care!”
He stormed out of the hall, slamming the heavy wooden door behind him. My
heart sank. I didn’t want to lose my brother, but even more than that, I didn’t
want to be alpha.
 3
NATALIE

“S eriously, Jorek, you should have heard them go on and on about how,
deep down, I really want a fated mate.”
I reached across the tiny table I shared with Jorek, my co-researcher and…
friend, for lack of a better word. Sienna and Arlynn were sure that he and I
were destined to be together, thanks to some freaky genetic quirk, but I
wasn’t falling for it.
I liked him, that much I couldn’t deny, but I wasn’t in love with him or
anything. Even when my arm brushed against his naked torso — Warg males
were almost always half naked, much to Arlynn’s delight — neither of us
flinched. But every time we got started discussing the tribe’s ‘drought’ of girl
babies, we barely knew the world around us existed.
“They’re only whelps, Natalie,” he said, handing me the herb sample I was
trying to grab. “They meant no offense.”
“I know, I know, it’s just so absurd.”
“What is?” He swiped a wild strand of hair that fell across his soft brown
eyes when looked up from the prehistoric piece of equipment he called a
microscope. I’d spent my entire life in labs and had never seen anything so
primitive. Salvage from some ancient shipwreck, he said. It was the best they
had here in the forest.
“Fated mates, of course. I get why children believe in it, but my intelligent,
worldly sisters?”
“So you still don’t believe such a thing is possible? That perhaps it’s in our
genetic makeup?”
I raised a skeptical eyebrow at Jorek as I prepped my own sample. “Oh,
you’ve found the gene that causes this magic, have you? Yeah, I didn’t think
so.”
He shrugged and returned to the microscope. “Just because we can’t see it—“
he shot a glance at me “—yet, doesn’t mean it doesn’t exist. Even if it
doesn’t, it’s so ingrained in our culture that to deny it is to deny our primary
paradigm.”
“Are you really telling me that my saying I don’t believe in fated mates
makes you question your very existence?”
A smirk tilted his lips as he studied his slide. “No. I know what we are, and
you will too, as soon as you accept the transformational bite.”
It was my turn to shrug.
“Honestly, it seems like cheating. I want someone to love me for who I am,
not because some random gene or bacteria or virus tells them to. And at the
rate we’re going at solving the riddle of why your people have trouble
conceiving females, I’ll have plenty of time to fall in love several times
before I become a Warg.”
“You’re really going to wait until we figure it out?”
“Absolutely. We’re on a roll, don’t you think, Jorek? I don’t want to mess up
my brain chemistry when we’re so close.”
He pulled the slide from the microscope with a sigh. “We’re not as close as
we thought. This isn’t it either. And you have the last sample.”
I slid the chipped glass slide under the lens, taking extra care because Jorek
only had a few of them. As I rotated the focus dial on the device, green and
yellow blobs came into sharp focus. Keeping my eyes on the blobs as best I
could, I added a drop of our control substance — an extract of a grain called
reet, which we were certain was one half of the equation — and held my
breath.
Tiny tan dots swirled around the blobs, spinning and zipping like crazy. This
was the most activity I’d seen with any sample of the flora found on the
Valley side of the river. My heart raced as fast as the dots. Could this be the
missing element we needed? If so, we’d just solved a generations-old
mystery.
The dots slowed, then stopped, then turned black. The breath I’d been
holding whooshed out of me in a curse.
“Shit!”
Jorek sighed. I leaned back in my chair and grabbed fistfuls of my hair.
Blonde strands stuck to my sweaty fingers. So close! I thought as I wiped my
hands clean.
“What now?” I asked.
“I have no idea. You know as much as I do, if not more, about the plants that
grow in the Valley. Are there any we’ve missed?”
“None. We even tried quadrapede silk.”
I tugged at the diaphanous material Wargs used as clothing. We were fairly
sure the second substance came from a plant, but I’d been getting desperate
by the time I tested it.
We sat in silence for a moment, deep in thought. When I’d heard about their
‘drought’, I immediately offered my services as a trained scientist. Not only
did I want to repay their kindness and generosity in taking us in when we so
desperately needed it, but I lusted for an intellectual challenge like this.
Jorek had isolated the reet compound before I arrived in the village. Together,
we theorized that the reet had to be mixed with some other plant to create a
completely new compound that promoted gender-balanced reproduction. Yet
none of the plants in the Valley performed as we’d hoped.
The silver thread of an idea wriggled around in my brain like a kronkworm,
always slipping just out of reach. The thread grew thicker and brighter until it
exploded in my brain like a supernova.
“Of course!” I shouted, jumping up. My chair flew across the lab — really
just one of many small huts in the village — but I paid it no mind. I barely
even registered the surprise on Jorek’s face.
“We’re idiots, Jorek! How could we not have seen this?”
“What?” He latched onto my excitement and jumped up, ready to be amazed.
I was about to blow his mind.
“We’ve only been testing plants from the Valley.” I stared at him expectantly.
Surely he’d get it. But he just stared back, waiting.
“So?”
“So we’ve been looking on the wrong side of the river. I guarantee you that
the mystery plant only grows over there.”
A crease formed between his thick eyebrows. “Why?”
“Think about it. When did this drought start? Generations before yours,
right?”
“Yes. I believe it was only a generation or two after the Great Split.”
“Exactly. Once Tooibas and Vanter had their little pissing match over a
woman and the Great Tribe split, boundaries were drawn. Which means
resources were split, too. The Valley Warg got the reet because it only grows
in the fertile soil down here.”
Jorek’s eyes lit up. “So there must be plants that only grow on the hill.”
“Right!”
I started jumping around the little hut, dancing with joy. Jorek followed for a
moment, and then stopped, clouds forming in his eyes. I stopped, breathless
and grinning.
“What?”
“How will we find it? Thrane can’t be happy with us right now.”
He had a point. Solan had pretty much just kicked his ass across the galaxy.
A search party for some random, unidentified plant probably would be seen
as an act of aggression or something. But if only one person went…
“I’ll go,” I said. “His sentries will have a hard time spotting just one person.
Besides, the tribe needs you more than me, so if anything goes wrong, you
can continue our work.”
Jorek took a step closer, gazing at me with wonder. “You’re truly
exceptional, Natalie. Is there anything you can’t do?”
A blush crept up my cheeks at his compliment. I wasn’t used to them. Back
in the city, and especially at the Training Center, I’d been shown every day
just how unimpressive I was. No one had ever given me respect, even my
teachers — as was proven when they sold me off as a sex slave instead of
sending me on a science-based mission, as promised.
“That settles it then,” I said, breaking eye contact and grabbing a few pieces
of equipment I’d need in the field. “I’ll set out right away.”
“Set out where?” asked a smooth voice behind me. I spun around to find
Chorn, Solan’s second-in-command leaning against the doorway. The
stealthy warrior could sneak up on a ghost.
“Oh, I’m just going to pop across the river and collect some different
samples. We’re so close, Chorn—“
“Excuse me?” he said, taking a step closer, a scowl rippling across his face.
“Across the river? Into the Hill tribe’s territory?”
I felt like a student being reprimanded by a mean teacher. God knows I had
plenty of those over the years.
“Yes?” I squeaked.
He barked out a scornful laugh. “A Terran female is simply going to walk
through our forest and not get eaten by any number of deadly creatures. Then
she’s going to somehow cross a raging river without drowning. And then—“
he laughed again “—and then she’s going to wander around Thrane’s lands
and not a single Hill Warg will notice her presence. Oh, and then return in
perfect health. Is that what you’re telling me?”
Perhaps I hadn’t thought it all through, but he didn’t need to be so rude about
it. Rather than give him the satisfaction of telling him he was right, I squared
my shoulders and sniffed at him.
“You have no idea what I’m capable of.” I glanced at Jorek. “Right, Jorek?”
We stood there, nose-to-nose and glaring at each other, waiting for Jorek’s
reply. When he stayed silent, I looked over at him again. His hair hung in his
face, his head dipped so I couldn’t catch his gaze.
Really?
When I turned back to Chorn, his expression had softened, but was no less
firm. “I’m sorry, Natalie. Your intentions are noble, but you wouldn’t make it
as far as the border.”
“But you don’t understand—“
“No, you don’t understand. You’re incapable of protecting yourself here.
Whatever is so important will just have to wait until Solan unites the tribes.”
He spun around on his heel and left me standing there, red-faced and fuming.
“Natalie, I’m s—“
I didn’t wait for Jorek to finish before stomping out of the lab.
 4
MARKON

I knocked lightly on Thrane’s door. He didn’t respond, but I heard him

moving around so I cautiously poked my head in. He stood over his bed,
stuffing a few possessions into a cloth sack.
“What are you doing?” I asked, closing the door behind me so no one could
overhear us.
“What does it look like?”
“It looks like you’re packing.”
“See, I knew you could figure it out,” he sneered. “You’ll make a terrific
alpha.”
A pit the size of a boulder settled in my gut.
“You’re alpha of the Hill Warg, Thrane. You always have been and you
always will be. You were born to it.”
Jamming a curved knife — thankfully still sheathed — into the sack, he
whirled on me. Fire danced in his orange eyes and it looked as if he’d ripped
out handfuls of hair.
“Don’t give me that, brother!” He jabbed a finger into my chest. “Don’t think
I didn’t notice your absence at the battle with the Valley scum. I didn’t say
anything before but now… You probably hoped Solan would kill me, didn’t
you? You’ve always been jealous of me!”
I rocked backward in shock that he’d think that of me. Of all the cruel jibes
Thrane had thrown at me, this hurt the most.
“I’ve never been jealous of you, Thrane. I’ve always looked up to you,
admired you. Since Father died, I’ve watched you handle the pressures of the
position with a confidence that astounds me.”
He shot me a skeptical glare. “Oh, so you’ve never once thought about
becoming alpha?”
I shrugged. “Of course, I did — when I was a whelp. But then I grew up and
realized that you’re the strong one. A fierce protector. A true leader. We need
that right now, Thrane. We need you.”
Thrane sighed and slumped to the bed, defeated. “No one needs me, little
brother. They don’t respect me anymore, and that’s one thing every good
leader must have.”
I sat next to him and nudged him with my shoulder. “I respect you.”
A hint of a smirk twitched his lips. “So I’ll lead a tribe of one?”
He shook his head and swiped a hand through his unruly hair, exposing his
flaming red scar again. He caught my gaze and my throat nearly closed at the
pain in his eyes.
“Why didn’t he just kill me, Markon? Why did Solan leave me alive? There’s
no honor in losing Holmgang, but at least you knew your opponent wouldn’t
force you to suffer the humiliation of surviving. He couldn’t even give me
that. Why?”
I’d never heard Thrane so despondent. All my life, he’d been my big brother,
always watching out for me. First, he protected me, and then he protected our
entire tribe. It was only natural that some Wargs chaffed under his strict
military-like command, but it was rare for us to lose one of our members so
they held their tongue and followed him faithfully.
But he was right. No one would look him in the eye any longer. If we walked
through the village together, they’d greet me but not him. It was painful to
watch the fall of my hero. I couldn’t imagine how rejected he felt.
“Remember the time you pulled me from the frothing maw of a hopwart?”
He gave me a ghost of a smile. “Father had punished you for stealing a jar of
dragonberry nectar from Mother so you ran away. Said you were going to
start your own tribe on the other side of the Hill. I have to admit, I was
flattered that you invited me to become the first member.”
“You told me that your place was here, with our tribe. You had a duty as the
first-born to take over after Father passed to the Great Beyond. But you
promised that when you were alpha, our tribes would never be at war. That
we would never turn out like Tooibas and Vanter.”
Thrane’s laugh wasn’t as robust as usual, but I was glad to hear it. “Right,
then you took off. Didn’t matter that you were only a whelp, and that even a
zeze fly could probably have carried you away.”
“That stinking hopwart got me instead. You know, I’ve never told you this,
but I’d never heard your beast roar so loudly before. Of course the thing that
really impressed me was when you took its head off with a single swipe of
your paw.”
It was Thrane’s turn to nudge me.
“Hey, it had my baby brother in its teeth. I was pissed!”
We laughed till tears dampened our eyes. We hadn’t had such an easy talk in
a long while. I missed it. When we caught our breath, I looked over at him.
“I needed you that day, Thrane. And I need you now. We need you.”
He sighed and shook his head. “Need and want are two different things,
Markon. Right now, it’s very clear that I’m not wanted.”
“That’s not true. The council agreed to keep you as alpha.” This caught his
attention.
“They did? What’s the catch?”
My eyes darted off the left as I tried to find the words. I was the more
diplomatic of the two of us, but even I had no idea how to break the news to
him.
“Nothing big, really. Only that I take some of the load off your shoulders.”
“How much of a load?” he asked through clenched teeth.
“Just the political stuff,” I hedged.
Thrane’s eyes narrowed to slits. “Such as this insane idea to merge with the
Valley scum?”
“That’s part of it.”
Thrane huffed and stood. He grabbed his bag and slung it over his shoulder.
“I won’t stay where I’m not wanted, Markon. The worst part of it is that I
can’t blame them for no longer respecting me. I know I wouldn’t.”
I stood to stop him, but he easily shrugged off my hand and move toward the
door. He stopped at the last minute and hope burst inside me that he’d
changed his mind. When he turned around, though, the blackness in his
expression drove a knife into my heart. His voice was a harsh whisper.
“Watch yourself, brother. There’s a spy in our midst. Don’t trust anyone.
Someone’s been feeding Solan information. He’s only trying to trick us into
lowering our defenses so he can attack and take our females. He can’t be
trusted.”
I let my eyes fall closed.
“No, he’s not, Thrane. Trust me.”
He barked his doubt.
“How would you know?”
I swallowed hard and steeled my nerves. There was no other choice. I looked
deep into his eyes, hoping the words would come and that he would
understand.
“Because I’m the spy.”
 5
NATALIE

“A rlynn, can I ask you something?”

“Sure,” she said, tucking another strand of my hair into an intricate pleated
braid she was trying to learn.
She stood behind my chair in the little hut we shared. Sienna had lived here
with us for a short time before she moved in with Solan. We both missed her,
but we were thrilled she’d found someone to take care of her. She’d always
been our mother hen, so it was about time she got pampered a little.
“Since your transformation into a Warg, do you…think any differently than
when you were just a Terran?”
“Just?”
“You know what I mean.”
Arlynn fell quiet for a moment.
“I don’t think so. Not so much that I’ve noticed, anyway.”
“That doesn’t make any sense. A bite that essentially changes you from one
species to another must do something to your brain, don’t you think?”
Arlynn shrugged and tugged at another strand. I continued questioning her.
“For example, do you think you’re as creative as before?”
“Oh, absolutely. If not more. Have you seen the mural I’ve been working on
inside the meeting hall? I’ve never made anything so majestic before.”
It was turning out to be quite a beautiful scene. I tried to nod but her tugging
and tucking stopped me. It was turning out to be quite a beautiful scene.
“But isn’t that because you only had access to small painting pads at the
Center? You never had a chance to try painting on a grand scale.”
“That’s true,” she conceded, though her tone sounded doubtful.
“What about your language skills?”
“Skills language fine my are.” I could feel her smirking behind me.
“Very funny. Seriously.”
“Seriously, you complain about my nonstop jabbering all the time.”
“But the other day you had trouble with the name of a plant.”
“You mean when I forgot ‘strothornium’?’” She gave my hair an extra hard
yank. “Give me a break, Nat. It’s not like I instinctively know the names for
all the weird stuff in this forest. What’s this all about, anyway?”
“Nothing,” I lied.
The truth was that Jorek’s refusal to defend me still stung like a swarm of
nettle bees. I thought he believed in me, but he didn’t even try to stand up for
me. I’d submitted to Arlynn’s hair torture hoping it would take my mind off
the situation.
“Is it because Chorn wouldn’t let you go look for more plants in Hill
territory?”
“Yes,” I said, eager to latch onto a more logical reason for trying to distract
myself than that a guy I kind of liked hurt my feelings.
“He’s right, you know. I wouldn’t even try going to the river alone. I’m too
new to this Warg thing. I’m sure I’d die.”
I had to snicker. “Yeah, you’d be more likely to hug a monster than to kill it.”
She thwapped me on the head, but giggled too.
“Do you ever wish you weren’t a Warg, that Thrane hadn’t give you the
bite?”
Arlynn didn’t answer at first and I worried I’d hurt her feelings. I had a
tendency to blurt things out that others found painful.
“Not exactly,” she finally said. “I wish it had been voluntary, though, because
I can’t help feeling something’s…missing.”
“What?”
“No idea. But something must be wrong since I haven’t found my fated mate
yet.”
I reached back and squeezed one of her suddenly trembling hands. “You will.
Remember what Solan said, that most mates are bonded at the claiming.”
“I’m sure you’re right,” she whispered, obviously not at all sure.
“Do you feel any different now that you’re a Warg? Emotionally speaking?” I
probed, honestly curious and doing my best to move the conversation in a
different direction.
“I’d have to say yes. I definitely feel my emotions more. For example, the
other day when you stole my piece of reet bread? As a Terran, I would have
been irked, but now, I was actually angry. Not like I wanted to kill you or
anything, just mad.”
“That’s comforting,” I teased. “It is interesting, though. Is it the same for
other emotions?”
“Well, I didn’t love you as much after that meal, but generally speaking, I
think my senses and emotions are heightened. Just the other day, I was
thinking about my parents and I thought my heart would explode from the
love I felt.”
I coughed in surprise. Arlynn had a blind spot when it came to the parents
who dumped her at the Training Center, and no amount of logic would
convince her that they didn’t love her and miss her every day.
“Your turn,” she said, tipping my head down slightly to access the hair on the
nape of my neck. “Tell me about your parents.”
Another random document with
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definitely ascertained. It may be an assumption that time will prove to
be unwarranted that all the Leptomedusae pass through a
Calyptoblastic hydrosome stage.

Fam. Aequoreidae.—In this family the hydrosome stage is not

known except in the genus Polycanna, in which it resembles a
Campanulariid. The sense-organs of the Medusae are statocysts.
The radial canals are very numerous, and the genital glands are in
the form of ropes of cells extending along the whole of their oral
surfaces. Aequorea is a fairly common genus, with a flattened
umbrella and a very rudimentary manubrium, which may attain a size
of 40 mm. in diameter.

Fam. Thaumantiidae.—The Medusae of this family are

distinguished from the Aequoreidae by having marginal ocelli in
place of statocysts. The hydrosome of Thaumantias alone is known,
and this is very similar to an Obelia.

Fam. Cannotidae.—The hydrosome is quite unknown. The

Medusae are ocellate, but the radial canals, instead of being
undivided, as in the Thaumantiidae, are four in number, and very
much ramified before reaching the ring canal. The tentacles are very
numerous. In the genus Polyorchis, from the Pacific coast of North
America, the four radial canals give rise to numerous lateral short
blind branches, and have therefore a remarkable pinnate
appearance.

Fam. Sertulariidae.—In this family the hydrothecae are sessile, and

arranged bilaterally on the stem and branches. The general form of
the colony is pinnate, the branches being usually on opposite sides
of the main stem. The gonophores are adelocodonic. Sertularia
forms more or less arborescent colonies, springing from a creeping
stolon attached to stones and shells. There are many species,
several of which are very common upon the British coast. Many
specimens are torn from their attachments by storms or by the trawls
of fishermen and cast up on the sand or beach with other zoophytes.
The popular name for one of the commonest species (S. abietina) is
the "sea-fir." The genus has a wide geographical and bathymetrical
range. Another common British species frequently thrown up by the
tide in great quantities is Hydrallmania falcata. It has slender spirally-
twisted stems and branches, and the hydrothecae are arranged
unilaterally.

The genus Grammaria, sometimes placed in a separate family, is

distinguished from Sertularia by several characters. The stem and
branches are composed of a number of tubes which are
considerably compressed. The genus is confined to the southern
seas.

Fam. Plumulariidae.—The hydrothecae are sessile, and arranged

in a single row on the stem and branches. Nematophores are always
present. Gonophores adelocodonic. This family is the largest and
most widely distributed of all the families of the Hydrozoa. Nutting
calculates that it contains more than one-fourth of all the Hydroids of
the world. Over 300 species have been described, and more than
half of these are found in the West Indian and Australian regions.
Representatives of the family occur in abundance in depths down to
300 fathoms, and not unfrequently to 500 fathoms. Only a few
species have occasionally been found in depths of over 1000
fathoms.

The presence of nematophores may be taken as the most

characteristic feature of the family, but similar structures are also
found in some species belonging to other families (p. 277).

The family is divided into two groups of genera, the Eleutheroplea

and the Statoplea. In the former the nematophores are mounted on
a slender pedicel, which admits of more or less movement, and in
the latter the nematophores are sessile. The genera Plumularia and
Antennularia belong to the Eleutheroplea. The former is a very large
genus, with several common British species, distinguished by the
terminal branches being pinnately disposed, and the latter,
represented by A. antennina and A. ramosa on the British coast, is
distinguished by the terminal branches being arranged in verticils.

The two most important genera of the Statoplea are Aglaophenia

and Cladocarpus. The former is represented by a few species in
European waters, the latter is only found in American waters.

Fam. Hydroceratinidae.—The colony consists of a mass of

entwined hydrorhiza, with a skeleton in the form of anastomosing
chitinous tubes. Hydrothecae scattered, tubular, and sessile.
Nematophores present. Gonophores probably adelocodonic.

This family was constituted for a remarkable hydroid, Clathrozoon

wilsoni, described by W. B. Spencer from Victoria.[317] The zooids
are sessile, and spring from more than one of the numerous
anastomosing tubes of the stem and branches. The whole of the
surface is studded with an enormous number of small and very
simple dactylozooids, protected by tubular nematophores. Only a
few specimens have hitherto been obtained, the largest being 10
inches in height by 4 inches in width. In general appearance it has
some resemblance to a dark coloured fan-shaped Gorgonia.

Fam. Campanulariidae.—The hydrothecae in this family are

pedunculate, and the gonophores adelocodonic.

In the cosmopolitan genus Campanularia the stem is monosiphonic,

and the hydrothecae bell-shaped. Several species of this genus are
very common in the rock pools of our coast between tide marks.
Halecium is characterised by the rudimentary character of its
hydrothecae, which are incapable of receiving the zooids even in
their maximum condition of retraction. The genus Lafoea is
remarkable for the development of a large number of tightly packed
gonothecae on the hydrorhiza, each of which contains a blastostyle,
bearing a single gonophore and, in the female, a single ovum. This
group of gonothecae was regarded as a distinct genus of Hydroids,
and was named Coppinia.[318] Lafoea dumosa with gonothecae of
the type described as Coppinia arcta occurs on the British coast.

Perisiphonia is an interesting genus from deep water off the Azores,

Australia, and New Zealand, with a stem composed of many distinct
tubes.

The genus Zygophylax, from 500 fathoms off the Cape Verde, is of
considerable interest in having a nematophore on each side of the
hydrotheca. According to Quelch it should be placed in a distinct
family.

Ophiodes has long and very active defensive zooids, protected by

nematophores. It is found in the Laminarian zone on the English
coast.

Fam. Eucopidae.—The hydrosome stage of this family is very

similar to that of the Campanulariidae, but the gonophores are free-
swimming Medusae of the Leptomedusan type.

One of the best-known genera is Obelia, of which several species

are among the commonest Hydroids of the British coast.

Clytia johnstoni is also a very common Hydroid, growing on red

algae or leaves of the weed Zostera. It consists of a number of
upright, simple, or slightly branched stems springing from a creeping
hydrorhiza. When liberated the Medusae are globular in form, with
four radial canals and four marginal tentacles, but this Medusa, like
many others of the order, undergoes considerable changes in form
before it reaches the sexually mature stage.

Phialidium temporarium is one of the commonest Medusae of our

coast, and sometimes occurs in shoals. It seems probable that it is
the Medusa of Clytia johnstoni.[319] By some authors the jelly-fish
known as Epenthesis is also believed to be the Medusa of a Clytia.

Fam. Dendrograptidae.—This family includes a number of fossils

which have certain distinct affinities with the Calyptoblastea. In
Dictyonema, common in the Ordovician rocks of Norway, but also
found in the Palaeozoic rocks of North America and elsewhere, the
fossil forms fan-shaped colonies of delicate filaments, united by
many transverse commissures, and in well-preserved specimens the
terminal branches bear well-marked uniserial hydrothecae. In some
species thecae of a different character, which have been interpreted
to be gonothecae and nematophores respectively, are found.

Other genera are Dendrograptus, Thamnograptus, and several

others from Silurian strata.

Order V. Graptolitoidea.
A large number of fossils, usually called Graptolites, occurring in
Palaeozoic strata, are generally regarded as the skeletal remains of
an ancient group of Hydrozoa.

In the simpler forms the fossil consists of a delicate straight rod

bearing on one side a series of small cups. It is suggested that the
cups contained hydroid zooids, and should therefore be regarded as
the equivalent of the hydrothecae, and that the axis represents the
axis of the colony or of a branch of the Calyptoblastea. In some of
the forms with two rows of cups on the axis (Diplograptus), however,
it has been shown that the cups are absent from a considerable
portion of one end of the axis, and that the axes of several radially
arranged individuals are fused together and united to a central
circular plate. Moreover, there is found in many specimens a series
of vesicles, a little larger in size than the cups, attached to the plate
and arranged in a circle at the base of the axes. These vesicles are
called the gonothecae.
The discovery of the central plate and of the so-called gonothecae
suggests that the usual comparison of a Graptolite with a Sertularian
Hydroid is erroneous, and that the colony or individual, when alive,
was a more or less radially symmetrical floating form, like a Medusa,
of which only the distal appendages (possibly tentacles) are
commonly preserved as fossils.

The evidence that the Graptolites were Hydrozoa is in reality very

slight, but the proof of their relationship to any other phylum of the
animal kingdom does not exist.[320] It is therefore convenient to
consider them in this place, and to regard them, provisionally, as
related to the Calyptoblastea.

The order is divided into three families.

Fam. 1. Monoprionidae.—Cups arranged uniserially on one side of

the axis.

The principal genera are Monograptus, with the axis straight, curved,
or helicoid, from many horizons in the Silurian strata; Rastrites, with
a spirally coiled axis, Silurian; Didymograptus, Ordovician; and
Coenograptus, Ordovician.

Fam. 2. Diprionidae.—Cups arranged in two or four vertical rows on

the axis.

Diplograptus, Ordovician and Silurian; Climacograptus, Ordovician

and Silurian; and Phyllograptus, in which the axis and cups are
arranged in such a manner that they resemble an ovate leaf.

Fam. 3. Retiolitidae.—Cups arranged biserially on a reticulate axis.

Retiolites, Ordovician and Silurian; Stomatograptus, Retiograptus,

and Glossograptus, Ordovician.
 Fossil Corals possibly allied to Hydrozoa.
Among the many fossil corals that are usually classified with the
Hydrozoa the genus Porosphaera is of interest as it is often
supposed to be related to Millepora. It consists of globular masses
about 10-20 mm. in diameter occurring in the Upper Cretaceous
strata. In the centre there is usually a foreign body around which the
coral was formed by concentric encrusting growth. Running radially
from pores on the surface to the centre, there are numerous tubules
which have a certain general resemblance to the pore-tubes of
Millepora. The monomorphic character of these tubes, their very
minute size, the absence of ampullae, and the general texture of the
corallum, are characters which separate this fossil very distinctly
from any recent Hydroid corals. Porosphaera, therefore, was
probably not a Hydrozoon, and certainly not related to the recent
Millepora.

Closely related to Porosphaera apparently are other globular,

ellipsoidal, or fusiform corals from various strata, such as Loftusia
from the Eocene of Persia, Parkeria from the Cambridge Greensand,
and Heterastridium from the Alpine Trias. In the last named there is
apparently a dimorphism of the radial tubes.

Allied to these genera, again, but occurring in the form of thick,

concentric, calcareous lamellae, are the genera Ellipsactinia and
Sphaeractinia from the Upper Jurassic.

Another important series of fossil corals is that of the family

Stromatoporidae. These fossils are found in great beds of immense
extent in many of the Palaeozoic rocks, and must have played an
important part in the geological processes of that period. They
consist of a series of calcareous lamellae, separated by considerable
intervals, encrusting foreign bodies of various kinds. Sometimes they
are flat and plate-like, sometimes globular or nodular in form. The
lamellae are in some cases perforated by tabulate, vertical, or radial
pores, but in many others these pores are absent. The zoological
position of the Stromatoporidae is very uncertain, but there is not at
present any very conclusive evidence that they are Hydrozoa.

Stromatopora is common in Devonian and also occurs in Silurian

strata. Cannopora from the Devonian has well-marked tabulate
pores, and is often found associated commensally with another coral
(Aulopora or Syringopora).

Order VI. Stylasterina.

The genera included in this order resemble Millepora in producing a
massive calcareous skeleton, and in showing a consistent
dimorphism of the zooids, but in many respects they exhibit great
divergence from the characters of the Milleporina.

The colony is arborescent in growth, the branches arising frequently

only in one plane, forming a flabellum. The calcareous skeleton is
perforated to a considerable depth by the gastrozooids,
dactylozooids, and nutritive canals, and the gastropores and
dactylopores are not provided with tabulae except in the genera
Pliobothrus and Sporadopora. The character which gives the order
its name is a conical, sometimes torch-like projection at the base of
the gastropore, called the "style," which carries a fold of the
ectoderm and endoderm layers of the body-wall, and may serve to
increase the absorptive surface of the digestive cavity. In some
genera a style is also present in the dactylopore, in which case it
serves as an additional surface for the attachment of the retractor
muscles. The pores are scattered on all aspects of the coral in the
genera Sporadopora, Errina, and Pliobothrus; in Spinipora and
Steganopora the scattered dactylopores are situated at the
extremities of tubular spines which project from the general surface
of the coral, the gastropores being situated irregularly between the
spines. In Phalangopora the pores are arranged in regular
longitudinal lines, and in Distichopora they are mainly in rows on the
edges of the flattened branches, a single row of gastropores being
flanked by a single row of dactylopores on each side. In the
remaining genera the pores are arranged in definite cycles, which
are frequently separated from one another by considerable intervals,
and have, particularly in the dried skeleton, a certain resemblance to
the calices of some of the Zoantharian corals.

In Cryptohelia the cycles are covered by a lid-like projection from the

neighbouring coenenchym (Fig. 136, l 1, l 2). The gastrozooids are
short, and are usually provided with a variable number of small
capitate tentacles. The dactylozooids are filiform and devoid of
tentacles, the endoderm of their axes being solid and scalariform.

The gonophores of the Stylasterina are situated in large oval or

spherical cavities called the ampullae, and their presence can
generally be detected by the dome-shaped projections they form on
the surface of the coral. The female gonophore consists of a saucer-
shaped pad of folded endoderm called the "trophodisc," which
serves the purpose of nourishing the single large yolk-laden egg it
bears; and a thin enveloping membrane composed of at least two
layers of cells. The egg is fertilised while it is still within the ampulla,
and does not escape to the exterior until it has reached the stage of
a solid ciliated larva. All the Stylasterina are therefore viviparous.
The male gonophore has a very much smaller trophodisc, which is
sometimes (Allopora) prolonged into a columnar process or spadix,
penetrating the greater part of the gonad. The spermatozoa escape
through a peculiar spout-like duct which perforates the superficial
wall of the ampulla. In some genera (Distichopora) there are several
male gonophores in each ampulla.

The gonophores of the Stylasterina have been regarded as much

altered medusiform gonophores, and this view may possibly prove to
be correct. At present, however, the evidence of their derivation from
Medusae is not conclusive, and it is possible that they may have had
a totally independent origin.

Distichopora and some species of Stylaster are found in shallow

water in the tropics, but most of the genera are confined to deep or
very deep water, and have a wide geographical distribution. No
species have been found hitherto within the British area.

Fig. 136.—A portion of a branch of Cryptohelia ramosa, showing the lids l 1 and l
2 covering the cyclosystems, the swellings produced by the ampullae in the
lids amp1, amp2, and the dactylozooids, dac. × 22. (After Hickson and
England.)

A few specimens of a species of Stylaster have been found in

Tertiary deposits and in some raised beaches of more recent origin,
but the order is not represented in the older strata.

Fam. Stylasteridae.—All the genera at present known are included

in this family.

Sporadopora is the only genus that presents a superficial general

resemblance to Millepora. It forms massive, branching white coralla,
with the pores scattered irregularly on the surface, and, like many
varieties of Millepora, not arranged in cyclosystems. It may, however,
be distinguished at once by the presence of a long, brush-like style
in each of the gastropores. The ampullae are large, but are usually
so deep-seated in the coenenchym that their presence cannot be
detected from the surface. It was found off the Rio de la Plata in 600
fathoms of water by the "Challenger."

In Errina the pores are sometimes irregularly scattered, but in E.

glabra they are arranged in rows on the sides of the branches, while
in E. ramosa the gastropores occur at the angles of the branches
only. The dactylopores are situated on nariform projections of the
corallum. The ampullae are prominent. There are several
gonophores in each ampulla of the male, but only one in each
ampulla of the female. This genus is very widely distributed in water
from 100 to 500 fathoms in depth.

Phalangopora differs from Errina in the absence of a style in the

gastropore; Mauritius.—Pliobothrus has also no style in the
gastropore, and is found in 100-600 fathoms of water off the
American Atlantic shores.

Distichopora is an important genus, which is found in nearly all the

shallow seas of the tropical and semi-tropical parts of the world, and
may even flourish in rock pools between tide marks. It is nearly
always brightly coloured—purple, violet, pale brown, or rose red. The
colony usually forms a small flabellum, with anastomosing branches,
and the pores are arranged in three rows, a middle row of
gastropores and two lateral rows of dactylopores on the sides of the
branches. There is a long style in each gastropore. The ampullae are
numerous and prominent, situated on the anterior and posterior
faces of the branches. Each ampulla contains a single gonophore in
the female colony and two or three gonophores in the male colony.

Spinipora is a rare genus from off the Rio de la Plata in 600 fathoms.
The branches are covered with blunt spines. These spines have a
short gutter-like groove at the apex, which leads into a dactylopore.
The gastropores are provided with a style and are situated between
the spines.

Steganopora[321] from the Djilolo Passage, in about 600 fathoms, is

very similar to Spinipora as regards external features, but differs
from it in the absence of styles in the gastropores, and in the wide
communications between the gastropores and dactylopores.

Stylaster is the largest and most widely distributed genus of the

family, and exhibits a considerable range of structure in the many
species it contains. It is found in all the warmer seas of the world,
living between tide marks at a few fathoms, and extending to depths
of 600 fathoms. Many specimens, but especially those from very
shallow water, are of a beautiful rose or pink colour. The corallum is
arborescent and usually flabelliform. The pores are distributed in
regular cyclosystems, sometimes on one face of the corallum only,
sometimes on the sides of the branches, and sometimes evenly
distributed. There are styles in both gastropores and dactylopores.

Allopora is difficult to separate from Stylaster, but the species are

usually more robust in habit, and the ampullae are not so prominent
as they are on the more delicate branches of Stylaster. It occurs at
depths of 100 fathoms in the Norwegian fjords. A very large red
species (A. nobilis) occurs in False Bay, Cape of Good Hope, in 30
fathoms of water. In this locality the coral occurs in great submarine
beds or forests, and the trawl that is passed over them is torn to
pieces by the hard, thick branches, some of which are an inch or
more in diameter.

Astylus is a genus found in the southern Philippine sea in 500

fathoms of water. It is distinguished from Stylaster by the absence of
a style in the gastropore.

Cryptohelia is an interesting genus found both in the Atlantic and

Pacific Oceans at depths of from 270 to about 600 fathoms. The
cyclosystems are covered by a projecting lid or operculum (Fig. 136,
l 1, l 2). There are no styles in either the gastropores or the
dactylopores. The ampullae are prominent, and are sometimes
situated in the lids. There are several gonophores in each ampulla of
the female colony, and a great many in the ampulla of the male
colony.

CHAPTER XI
 HYDROZOA (CONTINUED): TRACHOMEDUSAE—NARCOMEDUSAE—
SIPHONOPHORA

Order VII. Trachomedusae.

The orders Trachomedusae and Narcomedusae are probably closely
related to one another and to some of the families of Medusae at
present included in the order Calyptoblastea, and it seems probable
that when the life-histories of a few more genera are made known
the three orders will be united into one. Very little is known of the
hydrosome stage of the Trachomedusae, but Brooks[322] has shown
that in Liriope, and Murbach[323] that in Gonionema, the fertilised
ovum gives rise to a Hydra-like form, and in the latter this exhibits a
process of reproduction by gemmation before it gives rise to
Medusae. Any general statement, therefore, to the effect that the
development of the Trachomedusae is direct would be incorrect. The
fact that the hydrosomes already known are epizoic or free-
swimming does not afford a character of importance for distinction
from the Leptomedusae, for it is quite possible that in this order of
Medusae the hydrosomes of many genera may be similar in form
and habits to those of Liriope and Gonionema.

The free border of the umbrella of the Trachomedusae is entire; that

is to say, it is not lobed or fringed as it is in the Narcomedusae. The
sense-organs are statocysts, each consisting of a vesicle formed by
a more or less complete fold of the surrounding wall of the margin of
the umbrella, containing a reduced clapper-like tentacle loaded at its
extremity with a statolith.
 Fig. 137.—Liriope rosacea, one of the Geryoniidae, from the west side of North
and Central America. Size, 15-20 mm. Colour, rose. cp, Centripetal canal;
gon, gonad; M, mouth at the end of a long manubrium; ot, statocyst; t,
tentacle; to, tongue. (After Maas.)

This statocyst is innervated by the outer nerve ring. There appears to

be a very marked difference between these marginal sense-organs
in some of the best-known examples of Trachomedusae and the
corresponding organs of the Leptomedusae. The absence of a stalk
supporting the statolith and the innervation of the otocyst by the
inner instead of by the outer nerve ring in the Leptomedusae form
characters that may be of supplementary value, but cannot be
regarded as absolutely distinguishing the two orders. The statorhab
of the Trachomedusae is probably the more primitive of the two
types, and represents a marginal tentacle of the umbrella reduced in
size, loaded with a statolith and enclosed by the mesogloea.
Intermediate stages between this type and an ordinary tentacle have
already been discovered and described. In the type that is usually
found in the Leptomedusae the modified tentacle is still further
reduced, and all that can be recognised of it is the statolith attached
to the wall of the statocyst, but intermediate stages between the two
types are seen in the family Olindiidae, in which the stalk supporting
the statolith passes gradually into the tissue surrounding the statolith
on the one hand and the vesicle wall on the other. The radial canals
are four or eight in number or more numerous. They communicate at
the margin of the umbrella with a ring canal from which a number of
short blind tubes run in the umbrella-wall towards the centre of the
Medusa (Fig. 137, cp). These "centripetal canals" are subject to
considerable variation, but are useful characters in distinguishing the
Trachomedusae from the Leptomedusae. The tentacles are situated
on the margin of the umbrella, and are four or eight in number or, in
some cases, more numerous. The gonads are situated as in
Leptomedusae on the sub-umbrella aspect of the radial canals.

In Gonionema murbachii the fertilised eggs give rise to a free-

swimming ciliated larva of an oval shape with one pole longer and
narrower than the other. The mouth appears subsequently at the
narrower pole. The larva settles down upon the broader pole, the
mouth appears at the free extremity, and in a few days two, and later
two more, tentacles are formed (Fig. 138).

At this stage the larva may be said to be Hydra-like in character, and

as shown in Fig. 138 it feeds and lives an independent existence.
From its body-wall buds arise which separate from the parent and
give rise to similar Hydra-like individuals. An asexual generation thus
gives rise to new individuals by gemmation as in the hydrosome of
the Calyptoblastea. The origin of the Medusae from this Hydra-like
stage has not been satisfactorily determined, but it seems probable
that by a process of metamorphosis the hydriform persons are
directly changed into the Medusae.[324]

Fig. 138.—Hydra-like stage in the development of Gonionema murbachii. One of

the tentacles is carrying a worm (W) to the mouth. The tentacles are shown
very much contracted, but they are capable of extending to a length of 2
mm. Height of zooid about 1 mm. (After Perkins.)
In the development of Liriope the free-swimming larva develops into
a hydriform person with four tentacles and an enormously elongated
hypostome or manubrium; and, according to Brooks, it undergoes a
metamorphosis which directly converts it into a Medusa.

There can be very little doubt that in a large number of

Trachomedusae the development is direct, the fertilised ovum giving
rise to a medusome without the intervention of a hydrosome stage.
In some cases, however (Geryonia, etc.), the tentacles appear in
development before there is any trace of a sub-umbrella cavity, and
this has been interpreted to be a transitory but definite Hydroid
stage. It may be supposed that the elimination of the hydrosome
stage in these Coelenterates may be associated with their
adaptation to a life in the ocean far from the coast.

During the growth of the Medusa from the younger to the adult
stages several changes probably occur of a not unimportant
character, and it may prove that several genera now placed in the
same or even different families are stages in the development, of the
same species. In the development of Liriantha appendiculata,[325] for
example, four interradial tentacles appear in the first stage which
disappear and are replaced by four radial tentacles in the second
stage.

As with many other groups of free-swimming marine animals the

Trachomedusae have a very wide geographical distribution, and
some genera may prove to be almost cosmopolitan, but the majority
of the species appear to be characteristic of the warmer regions of
the high seas. Sometimes they are found at the surface, but more
usually they swim at a depth of a few fathoms to a hundred or more
from the surface. The Pectyllidae appear to be confined to the
bottom of the sea at great depths.

The principal families of the Trachomedusae are:—

Fam. Olindiidae.—This family appears to be structurally and in
development most closely related to the Leptomedusae, and is
indeed regarded by Goto[326] as closely related to the Eucopidae in
that order. They have two sets of tentacles, velar and exumbrellar;
the statocysts are numerous, two on each side of the exumbrellar
tentacles. Radial canals four or six. Manubrium well developed and
quadrate, with distinct lips. There is an adhesive disc on each
exumbrellar tentacle.

Genera: Olindias, Olindioides, Gonionema (Fig. 139), and Halicalyx.

As in other families of Medusae the distribution of the genera is very

wide. Olindias mülleri occurs in the Mediterranean, Olindioides
formosa off the coast of Japan, Gonionema murbachii is found in
abundance in the eel pond at Wood's Holl, United States of America,
and Halicalyx off Florida.

Two genera may be referred to in this place, although their

systematic position in relation to each other and to other Medusae
has not been satisfactorily determined.

Fig. 139.—Gonionema murbachii. Adult Medusa, shown inverted, and clinging to

the bottom. Nat. size. (After Perkins.)

Limnocodium sowerbyi is a small Medusa that was first discovered in

the Victoria regia tanks in the Botanic Gardens, Regent's Park,
London, in the year 1880. It has lately made its appearance in the
Victoria regia tank in the Parc de la Bête d'Or at Lyons.[327] As it
was, at the time of its discovery, the only fresh-water jelly-fish known,
it excited considerable interest, and this interest was not diminished
when the peculiarities of its structure were described by Lankester
and others. It has a rather flattened umbrella, with entire margin and
numerous marginal tentacles, the manubrium is long, quadrate, and
has four distinct lips. There are four radial canals, and the male
gonads (all the specimens discovered were of the male sex) are sac-
like bodies on the sub-umbrellar aspect of the middle points of the
four radial canals. In these characters the genus shows general
affinities with the Olindiidae. The difficult question of the origin of the
statoliths from the primary germ layers of the embryo and some
other points in the minute anatomy of the Medusa have suggested
the view that Limnocodium is not properly placed in any of the other
orders. Goto,[328] however, in a recent paper, confirms the view of
the affinities of Limnocodium with the Olindiidae.

The life-history of Limnocodium is not known, but a curious Hydroid

form attached to Pontederia roots was found in the same tank as the
Medusae, and this in all probability represents the hydrosome stage
of its development. The Medusae are formed apparently by a
process of transverse fission of the Hydroid stock[329] similar in
some respects to that observed in the production of certain
Acraspedote Medusae. This is quite unlike the asexual mode of
formation of Medusae in any other Craspedote form. The structure of
this hydrosome is, moreover, very different to that of any other
Hydroid, and consequently the relations of the genus with the
Trachomedusae cannot be regarded as very close.

Limnocodium has only been found in the somewhat artificial

conditions of the tanks in botanical gardens, and its native locality is
not known, but its association with the Victoria regia water-lily seems
to indicate that its home is in tropical South America.
Limnocnida tanganyicae is another remarkable fresh-water Medusa,
about seven-eights of an inch in diameter, found in the lakes
Tanganyika and Victoria Nyanza of Central Africa.[330] It differs from
Limnocodium in having a short collar-like manubrium with a large
round mouth two-thirds the diameter of the umbrella, and in several
other not unimportant particulars. It produces in May and June a
large number of Medusa-buds by gemmation on the manubrium, and
in August and September the sexual organs are formed in the same
situation.

Fig. 140.—Limnocnida tanganyicae. × 2. (After Günther.)

The fixed hydrosome stage, if such a stage occurs in the life-history,

has not been discovered; but Mr. Moore[331] believes that the
development is direct from ciliated planulae to the Medusae. The
occurrence of Limnocnida in Lake Tanganyika is supposed by the
same authority to afford a strong support to the view that this lake
represents the remnants of a sea which in Jurassic times spread
over part of the African continent. This theory has, however, been
adversely criticised from several sides.[332]

The character of the manubrium and the position of the sexual cells
suggest that Limnocnida has affinities with the Narcomedusae or
Anthomedusae, but the marginal sense-organs and the number and
position of the tentacles, showing considerable similarity with those
of Limnocodium, justify the more convenient plan of placing the two
genera in the same family.

Fam. Petasidae.—The genus Petasus is a small Medusa with four

radial canals, four gonads, four tentacles, and four free marginal
statorhabs. A few other genera associated with Petasus show simple
characters as regards the canals and the marginal organs, but as
very little is known of any of the genera the family may be regarded
as provisional only. Petasus is found in the Mediterranean and off
the Canaries.

Fam. Trachynemidae.—In this family there are eight radial canals,

and the statorhabs are sunk into a marginal vesicle. Trachynema,
characterised by its very long manubrium, is a not uncommon
Medusa of the Mediterranean and the eastern Atlantic Ocean. Many
of the species are small, but T. funerarium has sometimes a disc two
inches in diameter. Homoconema and Pentachogon have numerous
very short tentacles.

Fam. Pectyllidae.—This family contains a few deep-sea species

with characters similar to those of the preceding family, but the
tentacles are provided with terminal suckers. Pectyllis is found in the
Atlantic Ocean at depths of over 1000 fathoms.

Fam. Aglauridae.—The radial canals are eight in number and the

statorhabs are usually free. In the manubrium there is a rod-like
projection of the mesogloea from the aboral wall of the gastric cavity,
covered by a thin epithelium of endoderm, which occupies a
considerable portion of the lumen of the manubrium. This organ may
be called the tongue. Aglaura has an octagonal umbrella, and a
manubrium which does not project beyond the velum. It occurs in the
Atlantic Ocean and Mediterranean Sea.

Fam. Geryoniidae.—In this family there are four or six radial canals,
the statorhabs are sunk in the mesogloea, and a tongue is present in
the manubrium. Liriope (Fig. 137) is sometimes as much as three
inches in diameter. It has a very long manubrium, and the tongue
sometimes projects beyond the mouth. There are four very long
radial tentacles. It is found in the Atlantic Ocean, the Mediterranean
Sea, and the Pacific and Indian Oceans. Geryonia has a wider
geographical distribution than Liriope, and is sometimes four inches
in diameter. It differs from Liriope in having six, or a multiple of six,