How turtle shells evolved... twice Judy Cebra Thomas

Meet Odontochelys semitestacea.

This little creature spends its days
splashing in Late Triassic swamps

with a host of other reptiles.

Under the surface lies its best
defense against attack:

a hard shell on its belly.

Odontochelys is an early ancestor
of the turtle.

Its half-shelled body illustrates an
important point about the modern turtle:

it actually has two shells that develop
totally separately

while the turtle is still an embryo.

Both are extensions of the animal’s
skeleton,

and together they are made
of almost 60 bones.

Like other embryos,

turtle embryos are made of
undifferentiated cells

that become specific cell types,

and then organs and tissues,

through gene activity and communication
between cells.

At first, turtle embryos look very similar
to those of other reptiles,

birds, and mammals,

except for a bulge of cells called
the carapacial ridge.

The ridge expands around the body
between the neck and lower back,

creating a disc shape.

It guides the formation of the upper part
of the turtle’s shell,

called the carapace, likely by attracting
the cells that will become ribs.

Instead of curving downwards
to make a regular rib cage,

the ribs move outwards towards the
carapacial ridge.

They then secrete a signaling protein

that converts surrounding cells
into bone-forming cells.

These fifty bones grow until they meet
and connect with sutures.

A ring of bone solidifies
the carapace’s edges.

The outer layer of skin cells produces
the scales, known as scutes,

that cover the carapace.

The development of the bottom half
of the shell, the plastron,

is driven by neural crest cells,

which can produce a variety of different
cell types including neurons,

cartilage and bone.

A thick shield of these cells
spreads across the belly,

coming together in regions that produce
nine plate-like bones.

Eventually, these connect to the
carapace by sutures.

A turtle’s shell has obvious advantages
for guarding against predators,

but the rigid casing also presents
some challenges.

As the turtle grows,

the sutures between the bones
of the carapace and plastron spread.

Most mammals and reptiles rely on a
flexible rib cage

that expands to allow them to breathe,

but turtles use abdominal muscles
attached to the shell instead:

one to breathe in,
and one to breathe out.

So how did the shell evolve?

Though there are still gaps in the
fossil record,

the first step seems to have been
a thickening of the ribs.

The oldest known turtle ancestor,

a creature called Eunotosaurus africanus,

lived 260 million years ago and looked
almost nothing like a modern turtle,

but it had a set of broad, flat ribs

that anchored the muscles
of its powerful forearms.

Eunotosaurus was likely
a burrowing creature,

digging homes for itself in what’s
now southern Africa.

Odontochelys semitestacea illustrates
another, later step in turtle evolution,

with thick ribs like Eunotosaurus
plus a belly plate for protection.

Our first fossil evidence of the full
shell characteristic of modern turtles

is about 210 million years old,

and belongs to a species called
Proganochelys quenstedti,

whose ribs had fused.

Proganochelys could move
between water and land.

Unlike modern turtles, it couldn’t retract
its head into its shell,

but had defensive spines on its neck.

Modern turtle shells are almost as diverse
as the turtles themselves.

Sea turtles have flatter, lighter shells
for streamlined gliding through the water.

Land-dwelling tortoises, meanwhile,

have domed shells that can slip free of
predators’ jaws

and help them turn right-side up if
they fall on their backs.

Leatherback and softshell turtles

have shells without the ring of bone
around the edge of the carapace

or the tough scutes covering it,

making it easier for them to
squeeze into tight spaces.

认识 Odontochelys semitestacea。

这个小生物
在三叠纪晚期的沼泽中

与许多其他爬行动物一起嬉戏。

表面之下是它对
攻击的最佳防御:

它的腹部有一层坚硬的外壳。

Odontochelys 是龟的早期祖先

它半壳的身体
说明了现代龟的一个重要点:

它实际上有两个完全分开发育的壳,

而龟还是一个胚胎。

两者都是动物
骨骼的延伸

,它们一起
由近 60 块骨头组成。

像其他胚胎一样,

龟胚胎是由
未分化的细胞组成的

,这些细胞通过基因活动和细胞之间的交流,变成特定的细胞类型

,然后是器官和组织

起初,乌龟胚胎看起来
与其他爬行动物、

鸟类和哺乳动物的胚胎非常相似,

除了一个称为甲骨脊的细胞突起


在颈部和下背部之间的身体周围扩展,

形成一个圆盘形状。

它引导龟壳上部的形成

称为甲壳,可能是通过吸引
将成为肋骨的细胞。 肋骨

不是向下弯曲
以形成规则的肋骨,而是

向外朝向
甲骨脊移动。

然后它们分泌一种信号蛋白

,将周围的细胞
转化为骨形成细胞。

这五十块骨头一直生长,直到它们相遇
并与缝合线相连。

一圈骨头
使甲壳的边缘凝固。

皮肤细胞的外层产生覆盖甲壳
的鳞片,称为鳞片

。 壳

的下半部分
,即腹甲的发育

是由神经嵴细胞驱动的,神经嵴细胞

可以产生多种不同的
细胞类型,包括神经元、

软骨和骨骼。

这些细胞的厚盾
遍布腹部,

聚集在产生
九块板状骨骼的区域。

最终,这些
通过缝合线连接到甲壳。

龟壳在抵御捕食者方面具有明显的
优势,

但坚硬的外壳也带来了
一些挑战。

随着乌龟的成长,甲壳和腹甲

骨头之间的缝合线会
扩散。

大多数哺乳动物和爬行动物依靠一个
灵活的

胸腔扩张来呼吸,

但海龟使用
附着在壳上的腹肌来代替:

一个
吸气,一个呼气。

那么贝壳是如何进化的呢?

尽管化石记录仍然存在差距

,但第一步似乎
是肋骨的增厚。

已知最古老的

乌龟祖先,一种叫做非洲真龙的生物,

生活在 2.6 亿年前,看起来
几乎不像现代乌龟,

但它有一组宽而扁平的肋骨

,固定在
其强大的前臂肌肉上。

Eunotosaurus 很可能
是一种穴居动物,

在现在的南部非洲为自己挖掘家园

Odontochelys semitestacea 说明
了龟进化的另一个较晚的步骤,

具有像 Eunotosaurus 一样的厚肋骨
以及用于保护的腹板。

我们第一个
关于现代海龟全壳特征的化石

证据大约有 2.1 亿年的历史

,属于一种叫做
Proganochelys quenstedti 的物种,

它的肋骨已经融合。

Proganochelys 可以
在水和陆地之间移动。

与现代海龟不同,它不能
把头缩进壳里,

但脖子上有防御刺。

现代海龟壳几乎
与海龟本身一样多样化。

海龟有更扁平、更轻的
壳,可以在水中流线型滑行。

与此同时,陆生陆龟

有圆顶壳,可以从
捕食者的下颚

滑落,如果它们仰面倒地,可以帮助它们正面朝上。

棱皮龟和软壳龟

的壳没有
围绕甲壳边缘的骨环

或覆盖它的坚韧的盾甲,

使它们更容易
挤进狭窄的空间。