论文出处：
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https://royalsocietypublishing.org/rsif/article/23/237/20250523/481493
Scorpions diverged from their closest relatives around the Ordovician Period,
and since then, environmental interactions have shaped the evolution of the
material properties of their exoskeletons. Hardening of this structure via
the incorporation of transition metals has enabled biomechanical advancements
in weapon development.
蝎子大约在奥陶纪时期与其近亲分化，此后，环境相互作用塑造了其外骨骼材料特性的演
化。过渡金属的掺入增强了这种结构的硬度，从而促进了武器生物力学的发展。
Scorpion weapons consist of the stinger (telson) and claws (chelae) and
contain diverse metals such as zinc, manganese and iron, though little is
known about comparative patterns of incorporation across the wider clade.
蝎子的武器由尾刺（尾节）和螯肢组成，含有锌、锰和铁等多种金属，但人们对整个蝎子
进化分支中这些金属的掺入模式知之甚少。
In this study, we harness X-ray-driven microanalytical techniques to
characterize the different elemental enrichment patterns within the weapons
of 18 species from a range of scorpion families.
在本研究中，我们利用X射线驱动的微分析技术，对来自不同蝎子科的18个物种的武器中
不同的元素富集模式进行了检测。
We hypothesized that enrichment by metal would be inversely correlated
between weapons, tied to their functional roles and morphological diversity.
We identified cryptic enrichment strategies, including weapon-selective
elemental replacement and an inverse enrichment of Zn between weapons. Chela
enrichment by Zn was found to positively correlate with a morphological
indicator of chelae pinch strength, wherein Zn enrichment was greater in
specimens with reduced crushing power. This study supports a growing body of
research into the evolution of metal enrichment among invertebrates and
provides a greater understanding of the material properties of the
exoskeleton within weapon development.
我们假设不同武器的金属富集程度呈负相关，这与其功能性和形态多样性有关。我们发现
了一些隐藏的富集策略，包括武器选择性的元素替换以及武器间锌含量的负相关。研究发
现，螯肢锌的富集程度与螯肢的捏力形态指标呈正相关，即捏力越弱的样本，锌的富集程
度越高。这项研究为无脊椎动物金属富集演化的研究提供了支持，并加深了我们对武器研
发中外骨骼材料特性的理解。
The predominant incorporation of transition metals during sclerotization,
even in the earliest stages of life, has been shown as a driver of hardness
within the cuticle of chelicerates . Offering improved biomechanical
properties, the enhanced material allows for the development of sharper,
damage-resistant weapons such as fangs, aculei (needle-like envenoming
structure), ovipositors (tube structure to lay eggs) and chelae .
Termed ‘heavy element biomaterials’ or HEBs, zinc (Zn), manganese (Mn) and
iron (Fe) are the most frequently observed elements appearing in enriched
cuticle as metal–ligand coordinated bonds .
An association between HEBs and lighter elements, including calcium (Ca) and
chlorine (Cl), has also been reported .
即使在生命早期阶段，硬化过程中过渡金属的大量掺入也已被证实是螯肢动物角质层硬度
的主要驱动因素。这种增强的材料具有更优异的生物力学性能，使其能够发育出更
锋利、更耐损伤的武器，例如毒牙、刺突（针状毒刺结构）、产卵管（用于产卵的管状结
构）和螯肢。
锌（Zn）、锰（Mn）和铁（Fe）是富集角质层中最常见的元素，它们以金属-配体配位键
的形式存在，被称为“重元素生物材料”（HEBs）。
也有报告指出，HEBs与钙（Ca）和氯（Cl）等较轻元素存在关联。
Given the diversity of arthropods, with over a million species described and
an estimated 2–6 million total , there remains much to learn about
cuticular metal enrichment.
While metal enrichment has been demonstrated to
be widely observed across Arthropoda , study in this topic has only
occurred in a handful of species. Thus, we still know very little about how
it is evolving within and between different groups. It is likely that a lack
of study, as opposed to the absence of enrichment, is a determining factor in
this observation.
The ancient terrestrialization of Myriapoda, Hexapoda and
Arachnida between the Cambrian and Silurian Periods appears to be the
driving factor in the emergence of cuticular HEB enrichment in arthropods.
Alternatively, the majority of marine Mandibulata employ biomineralization by
the more abundant calcium, in the form of calcium carbonate.
Exceptions can be found, including metal enrichment in copepods ,
which are marine or aquatic, and a total absence of observable enrichment in
mites (Parasitoformes/Acariformes) .
Although current sampling to date has been limited compared with known total
organismal diversity, the presence of enrichment across distant lineages is
considered a strong example of convergent evolution and has led to a rapidly
growing field of interest .
鉴于节肢动物的多样性，已描述的物种超过一百万种，估计总数在200万至600万之间
，我们对表皮金属富集仍知之甚少。虽然金属富集已被证实广泛存在于节肢动物，但
对此的研究仅限于少数物种。
因此，我们对不同类群内部和类群之间金属富集的演化过程仍然知之甚少。研究的缺乏，
而非富集现象的缺失，很可能是造成此现象的决定性因素。
寒武纪至志留纪期间，多足纲、六足纲和蛛形纲的古老陆生化似乎是节肢
动物表皮高能生物富集（HEB）出现的主要驱动因素。另一方面，大多数海洋有颚类动物
则利用更丰富的钙（以碳酸钙的形式）进行生物矿化。
例外情况也存在，例如桡足类动物（海洋或水生生物）中金属富集以及螨类（寄螨目/
蜱螨目）中完全未观察到金属富集。
尽管迄今为止的采样与已知的生物多样性总量相比仍然有限，但远缘谱系中金属富集现象
被认为是趋同进化的一个有力例证，并已催生出一个迅速发展的研究领域。