真实存在于特兰西瓦尼亚的史上最怪异的恐龙

Most of us know of Transylvania, an historical region in Romania, from Bram Stoker's novel, Dracula. But even this fictional thriller almost pales in comparison to what really took place in Transylvania around 70 million years ago.

我们大部分人都从布拉姆·史托克的小说《德拉库拉伯爵》一书中了解了特兰西瓦尼亚这一罗马尼亚的历史古迹。但即使是这虚幻情节的惊悚程度与 7 千万年前在特兰西瓦尼亚所真正发生的事情相比也显得几乎是平淡无奇的了。

The linked cast of characters from then until now includes dwarfs, an eccentric Austro-Hungarian motorcycle-riding baron, and now a kickboxing dinosaur with off-the-chart claws.

与之有关的人物是矮人和一位怪异的骑摩托的奥匈帝国男爵，以及现在我们所知的拥有难以想象的爪子的跆拳道恐龙。

(Dwarf Dinosaurs; Credit: Vlad Codrea)

（矮恐龙：版权：Vlad Codrea）

(Baron von Nopsca; Wikimedia Image)

（Nopsca 男爵：维基百科图片）

(The Kickboxing Dinosaur: Balaur bondoc; Credit: Mick Ellison, Zoltan Csiki, Matyas Vremir, Stephan Brusatte, Mark Norell)

（跆拳道恐龙：巴拉乌尔·邦多克龙：版权：Mick Elison、Zoltan Csiki、Matyas Vremir、Stephan Brusatte、Mark Norell）

The motorcycle rider, Baron von Nopsca, comes into the real life story circa 1900. During that year, his sister found a tiny bone on the family estate in Transylvania. The baron, who was a dinosaur buff, identified the bone as belonging to a dwarf dinosaur that likely once lived on an island in the region.

摩托骑士 Nopsca 男爵的真实故事要追溯到大约 1900 年。在那一年，他的姐姐在特兰西瓦尼亚的家族庄园内发现了一块小骨头。这位恐龙迷男爵将这块骨头鉴定为一条可能曾经在此地区的岛屿上生存的矮恐龙。

His theory was immediately ridiculed and largely dismissed. But over a century later, paleontologists have proven that most of Baron von Nopsca's discoveries and theories were legit.

他的理论立刻受到了嘲笑，被主流忽视。不过在一个世纪以后，古生物学者们证明了 Nopsca 男爵的大部分发现和理论都是正确的。

Earlier this year, I wrote about new evidence further supporting the dwarf dinosaur claims. Today comes the news that a predatory dinosaur, the "kickboxer," likely feasted on the dwarfs and goodness knows what else.

今年初，我发表了更进一步支持矮恐龙学说的新证据。今天就有条新闻说一种叫做“跆拳道”的食肉恐龙很可能以矮人为食，其他的情况只有上帝才知道了。

Below is a transcript of my interview with University of Bucharest paleontologist Zoltan Csiki, who led the research project concerning Balaur bondoc:

下面是我和布加勒斯特大学的古生物学家佐尔坦·蒂克尔的采访抄录，他是有关巴拉乌尔·邦多克龙的研究项目的领头人：

Jennifer Viegas: Is it true that this dinosaur had two enormous claws on each of its feet? And isn't that unprecedented for a dromaeosaur?

珍妮佛·维埃加斯：这种恐龙是不是真的脚上各长着两只巨大的爪子？这对于厉爪属恐龙来说是不是从未见过的？

Zoltan Csiki: Yes, indeed, it had two large sickle claws on both feet. This is not only unlike any other dromaeosaurid known to date, with only one enlarged claw on each foot, but unlike almost all other theropods (the group of mainly predatory dinosaurs to which Balaur belongs), in which the reduction of the digits is the norm, having only three functional digits. Balaur is outstanding because it had both FOUR functional digits on its hind limb, including TWO enlarged and hyperextensive claws.

佐尔坦·蒂克尔：没错，就是这样，它的双脚各有着两个巨大的镰刀状的爪子。这不仅和目前已知的其他脚上有着一只大爪子的厉爪属恐龙不一样，而且也和大部分其他兽脚亚目食肉恐龙（这类恐龙主要是食肉的，巴拉乌尔龙也属于这类）不同，这类恐龙的进化方向是减少脚趾数量，只有三个活动的脚趾。巴拉乌尔龙突出的地方在于它的后肢拥有 2 个大号的不受关节限制自由活动的爪子，还都有着 4 个活动脚趾。

(Balaur bondoc's double sickle-clawed foot: Credit: Mick Ellison)

（巴拉乌尔·邦多克龙的两只镰刀状的脚：版权：Mick Elison）

JV: Your PNAS paper mentions that Balaur bondoc had large muscles around its pelvis area, as well as fused bones. How then would this dinosaur have moved?

珍：你在美国国家科学院院刊上的论文提到巴拉乌尔·邦多克龙的骨盆附近有着很大的肌肉，而且骨骼之间有着固定的连接。那么这种恐龙要如何移动呢？

ZC:

佐：

Large pelvic muscles mean large forces exerted along the hind limbs – it looks that Balaur had more powerful hind limbs than its relatives. However, strong and powerful legs do not necessarily mean speed; and indeed, while other dromaeosaurids such as Velociraptor and Deinonychus (both close relatives of Balaur) are usually depicted as fast runners, Balaur almost certainly was not one like that. Not a sprinter, Balaur was probably using his powerful hind limbs for a different function – but we are as yet unsure what that might have been. The foot, equipped with the double set of sickle claws, suggests it used these to attack prey; functional morphology studies and lucky discoveries such as the famous “fighting dinosaurs” from Mongolia allow us to infer that dromaeosaurids did use their enlarged claws to attack prey, and can hypothesize that Balaur used them the same way. But either it delivered its blows in a more powerful way, or else it might have used his powerful hip and tight musculature for other reasons, not necessarily directly related to its hunting strategy. Detailed functional anatomy studies are needed to clarify this issue, and this is planned for the future.

骨盆肌肉大就说明后腿施展的力量大——看来巴拉乌尔龙比它的近亲们有着更强的后腿力量。但是，强壮而有力的腿并不能断言奔跑速度的快慢。事实上，虽然例如速龙和恐爪龙（两者都是巴拉乌尔龙的近亲）这样的其它厉爪属恐龙通常被描述为恐龙中的赛跑冠军，但巴拉乌尔龙几乎完全不是这么回事儿。它不是短跑健将，巴拉乌尔龙可能使用它有力的后腿进行其他的行动——但我们目前还不确定这会是什么能力。它的脚有着两只镰刀状的爪子，表明它用它们来攻击猎物。功能形态学以及类似著名的蒙古利亚“战斗恐龙”的幸运发现让我们能推断有羽类恐龙确实是用它们的大爪子攻击猎物，并得以假设巴拉乌尔龙也是如此。但要么它以更有力的方式发动它的爪击，要么它可能在其他方面利用其强有力的臀部和固定连接的肌肉组织，我们并不能就此将它与其猎食策略直接联系起来。我们需要详细的功能解剖学研究来解开这个谜题，这也是我们将来要做的。

As for the fused hand and foot bones, we have no clear ideas as to their exact use. Fusion in the hind-limb is used sometimes to stabilize limb posture, especially along the most unstable ankle joint, if the animal lives and moves in mechanically stressful environments. On the other hand, we see similar modifications (even if not to the extent seen in Balaur) in relatively closely related taxa such as birds or Velociraptor. So whether these are phylogenetically or ecologically driven modifications, is unclear for the moment, and should be explained only after detailed functional morphology analyses of Balaur.

对于手足骨骼的固定连接的具体用途，我们还没有得出明确的结论。如果这种恐龙生存和活动的环境对机械运动的压力很大，那么后腿骨骼的固定连接有时可以用来稳固后腿的姿势，特别是在最不稳固的踝关节处。在另一方面，我们从类似鸟类或速龙类这样的密切相关的物种中可以找到类似的改变（不过没有巴拉乌尔龙这样的程度）。因此无论这是由进化还是生态所驱使的改变，目前还不确定，不过应该只要对巴拉乌尔龙经过详细的功能形态学分析后就会水落石出。

JV: Please describe a possible scenario in which this predatory dinosaur attacks, kills and consumes one of its prey species.

珍：请描述一下这种食肉恐龙攻击、捕杀并猎食某只猎物的场景。

ZC: As we do not have as yet a good understanding of the lifestyle and hunting habits of this animal, imaging such a scenario would be a little far-fetched. For the moment what is sure is that, if functional morphology reasoning is reliable to any reasonable degree, Balaur did what it did in a markedly different manner than its close relatives.

佐：因为我们目前还没有很好地了解这种动物的生活风格与猎食习惯，要想想象出这样一个场景就有点牵强附会了。目前我们所确定的是，如果功能形态学的结果在任何方面能值得信赖的话，巴拉乌尔龙的行为方式与它的近亲有着截然的不同。

JV: Why do you think this dinosaur evolved such unusual features? And why didn't it shrink to become a dwarf, as so many of the other dinosaurs on islands did?

珍：你为什么觉得这种恐龙进化出了如此不寻常的特点？而为什么它没有缩小为矮恐龙，就像岛上的许多其他恐龙一样呢？

ZC: Our most parsimonious explanation for the highly unusual (autapomorphic ) nature of Balaur is that it represents a by-product of its insular habitat. Animals tend to do strange things on islands, simply because they are allowed to – selective forces and restrains are more relaxed in island environments than within the more competitive continental environments. But even these “strange things” must have had biological explanations, being the results of different constrains related to its lifestyle, feeding strategies, social structure, sexual behavior and so on. Because Balaur is so unlike any other dinosaur for which a reasonable connection between morphology and function was suggested, for the moment we (or at least I) do not have a sound understanding of the nature of the evolutionary forces that drove its peculiar anatomical adaptations. But hopefully we’ll be able to better answer these questions in the future.

佐：我们对巴拉乌尔龙这种高度不寻常（在家族中独树一帜）的天然特征的最保守解释是它是岛屿栖息地所带来的副作用的表现。在岛上的动物会倾向于做些古怪的事情，这不过是因为他们可以这么做——选择性的压力和抑制在岛屿环境中比在竞争性更强的大陆环境中更加宽松。不过即使是这种“古怪的事情”也一定有着生物学上的解释，是与其生活风格、捕食策略、社会结构、性行为等等有关的不同限制所决定的。因为巴拉乌尔龙和其他的恐龙截然不同，可能是由于形态学和功能学的合理结合，目前我们（或者说至少我）没有一种合理的解释，究竟是什么样的自然力量驱使它进化产生了独特的解剖学环境适应结果。但我们很有信心将在未来对这些问题得出更好的解答。

What is important for the moment is that the discovery of Balaur brought us in front of the recognition that we are still far from fully comprehending the complexity and diversity of Mesozoic life, not even within groups so thoroughly studied and considered relatively “conservative” as the dromaeosaurids. It's something like the discovery of a lion with elephantine hind limbs and a grasping ape-like hand – it would do such unexpected and unsuspected things that these would be difficult (near to impossible) to be deduced by simply watching the daily activities of its closest relatives, the lions, which will feed, hunt, mate, and behave completely differently. Most probably so did Balaur as well, and this is the main challenge coming out of its discovery – understanding how it actually lived.

目前重要的事情是巴拉乌尔龙的发现让我们再次意识到我们依然远远没有完全理解中生代生物的复杂和多样性，甚至是在研究得非常透彻，被认为是相对较“传统”的厉爪属恐龙这样的种群中亦是如此。它就像是头有着巨大的后腿和如猿猴一样可以攒握的双手的狮子——它可能会做出超乎想象难以预计的事情，光靠观察其捕食、猎杀、交配和行为都与之完全不同的近亲狮子的日常行动很难（几乎不可能）推论出来。巴拉乌尔龙很可能也是这样，而这就是这一发现之后所带来的主要挑战——理解它实际的生活方式。

Size reduction, as an evolutionary consequence of island habitation, is not universal, it's not a “law” of nature; it is only a case made based on a statistical tendency for such a size change, observed in a wide range of animals, a “rule." And this rule states that, once entering an island, large animals tend to become smaller, generally because of space and resource limitations, while small ones might become larger. However, some modern predators (such as mammalian carnivores) were suggested to be excepted from this “rule”, as they do not regularly dwarf in island habitats. It is thus possible that the same was applicable to Mesozoic predators as well. On the other hand, the relatives of Balaur (the velociraptorines) were already relatively small in absolute size, so that possibly they were below a threshold size that would have triggered dwarfing, even on an island, possibly because it had sufficient food resources, or because its main prey species, although dwarfed, were still large enough to not be taken down by a severely downsized predator.

体格的缩小是一项岛屿栖息地的进化结果，它并非是通用的，它并非自然的“法则”。这不过是根据各种各样的动物的观察结果所得出的这类体格变化的统计规律“定律”罢了。而这一定律表明，一旦进入一个岛屿，大型动物就会倾向于变小，通常是由于空间和资源的限制，而小的物种可能会变大。但是，一些现代食肉动物（例如哺乳类食肉动物）被排除在这一定律之外，因为他们并没有在岛屿栖息地中如通常那样变小。于是中生代的食肉动物也可能是这样。在另一方面，巴拉乌尔龙的近亲（速龙）的体型已经相对很小了，因此他们可能没有达到需要变小的那个尺寸界线，即使在岛屿上也是如此，可能是由于它有着充足的食物资源，或者由于它的猎物虽然变小了，但依然足够大，不需要让捕食者大幅缩小来适应它。

(Balaur bondoc; Credit: Mick Ellison)

（巴拉乌尔·邦多克龙：版权：Mick Elison）

JV: Is it thought that land bridges connected this island (now Romania) to the mainland, helping to explain how dinosaurs came to the island in the first place?

珍：连接这个岛屿（现在的罗马尼亚）和大陆的陆地桥是否有助于解释这些恐龙是如何首先登上这个岛上的？

ZC: That Balaur is so closely related to animals living in the same time in other parts of the world (such as Saurornitholestes in North America or Velociraptor in Asia) clearly demonstrates that it is not the descendant of a long line of animals that would have lived in isolation in the Late Cretaceous archipelago – while this is indeed the case of other dinosaurs, turtles or mammals from the same assemblage. This means, in turn, that the line leading to Balaur separated relatively recently from the larger stock of velociraptorines (the likes of Velociraptor or Deinonychus, its closest relatives), more recently that the barriers (seaways) separating its homeland from other continental areas were put in place before its divergence, and that the immediate ancestors of Balaur must have moved somehow across these barriers. We do not yet have a definitive idea as to the ways this migration happened. Intermittent faunal connections do not necessarily require land bridges to connect isolated areas only to be afterwards destroyed; they can occur as results of “island hopping” (certainly a good candidate hypothesis in the case of the Late Cretaceous South European archipelago), as chance dispersal over marine barriers (we know that recent terrestrial animals can spread actively or be rafted passively across stretches of marine waters) or during sea-level drops that would have united previously isolated areas. Nor do we have an idea as to the timing of these faunal connections. Actually, it looks that they took place several times during the Late Cretaceous, allowing the dispersal of different groups of animals from and to Asia, North America or the southern Gondwanan landmasses (Africa, South America, India). A more sound phylogeny of Balaur, together with a better understanding of the geological evolution of southern Europe, is required to pinpoint the mode, timing and even the direction of these faunal connections.

佐：巴拉乌尔龙和同时期在世界其他地区生活的动物（例如北美的蜥鸟盗龙或者亚洲的速龙）的血缘关系非常密切，这明确说明了它并非晚白垩世生活在与世隔绝的岛屿上的很长的动物进化链的后裔——不过这确实是某些其他类似情况的恐龙、乌龟或者哺乳动物的状况。这反过来说明了指向巴拉乌尔龙的进化链从较为庞大的速龙类（速龙和其近亲恐爪龙之类的生物）分支分离出去的时间是较晚的，而将其家园与其他大陆区域分开的障碍（海峡）出现是在其进化分支之后的事情，而巴拉乌尔龙的直系祖先一定通过什么方式渡过了这些障碍。我们目前还不确定这一次迁移是如何发生的。间歇性的动物物种联系并不一定需要大陆桥来连接之后会被摧毁的隔离区域；它们可能是由于“岛屿跳跃”而产生的结果（这当然是可以适用于晚白垩世南欧岛屿状况的很好的候选假说），而可能偶然地通过海洋障碍传播（我们知道近代的陆地动物可以主动渡海或者被动地被木筏什么的带走读过长距离的海路）或者趁着落潮时暂时连接隔离区域的时机。我们也不知道这些动物物种联系的时间表。实际上，似乎在晚白垩世期间发生了许多次，让不同种群的动物来往分散于亚洲、北美或者南冈瓦纳陆块（非洲、南美、印度）。要得到更合理的巴拉乌尔龙的进化史，以及更好地理解南欧的地质演化，我们需要精确定位这些动物物种联系的方式、时间，甚至以及方向。

But, the bottomline is – Balaur being so closely related to animals from Asia and North America suggests interchanges with that area. Besides understanding the lifestyle and functional morphology of this peculiar animal, this is a second important challenge for us – to decipher its origin and ways of evolving.

但是，至少我们知道——巴拉乌尔龙和亚洲和北美的动物关系如此密切，表明这些区域之间有物种交换。除了理解这独特物种的生活风格和功能形态学，我们的次要挑战是——解译其来源和进化历程。

JV: How many millions of years ago did this predatory dinosaur live, and when did it die out?

珍：这种食肉恐龙生活在几百万年前，又是在什么时候灭绝的？

ZC: Dating of continental deposits, such as those yielding the remains of Balaur, are often difficult, as these usually lack good age indicators. Nevertheless, we can reliably restrain the age of these deposits (and thus that of Balaur) to the Maastrichtian, that is, the last stage of the Cretaceous, spanning the 72 to 65 Mya time period, because these continental deposits rest on top of marine ones which can be dated using different fossils as ending with the uppermost Campanian (the stage of the Upper Cretaceous preceeding the Maastrichtian). That means that our record of Balaur is restricted to the very end of the Cretaceous. But it is not impossible that it (or its ancestor species) might have lived during earlier time periods in the European archipelago.

佐：追溯例如让巴拉乌尔龙遭灭顶之灾的大陆沉降的时间往往困难重重，因为这些通常缺少较好的年代标识。然而，我们能有把握地将这些沉降（也包括巴拉乌尔龙的）的年代可能缩小到麦斯里希特阶，也就是白垩纪最后阶段，约在 72 至 65 Mya 时间段内，这是由于这些大陆沉降留下一些在海面上的部分中的不同化石可以追溯到坎帕阶（自晚白垩世至麦斯里希特阶这一阶段）。这意味着我们有关巴拉乌尔龙的记录限定于白垩纪的晚期的晚期。不过它（或其祖先物种）不可能在更早的时间在欧洲群岛生活。

On the other hand, there is no data about where it went extinct. Usually dinosaurs are considered to be one of the flagship casualties of the Cretaceous-Tertiary boundary events, and thus the most obvious answer would be – at the end of the Cretaceous. But, as we have only two specimens of Balaur known as yet, whose relative ages are unknown, we cannot say for sure whether it (or any of its contemporary dinosaur species) lived up to the very end of the Cretaceous or if eventually went extinct earlier. There is just not enough data to decide.

另一方面，没有任何有关其灭绝时刻的数据。通常我们认为恐龙是白垩纪——第三纪过渡过程中主要的灭绝物种，因此最明显的答案是——在白垩纪末期。不过，由于我们目前只有两个巴拉乌尔龙的样本，其相关年代还未知，因此我们不能确切地说它是否（或者其同代的恐龙物种中的任何一种）生存到了白垩纪的晚期的晚期，还是最终在更早的时候就已经灭绝。尚没有足够的数据来定论。

JV: In a nutshell, please explain why this dinosaur was so "aberrant" and "strange?"

珍：请你简单地解释一下这种恐龙为何如此“异常”和“奇怪”？

ZC: Balaur was aberrant and strange first of all because it is morphologically so divergent and shows so many unique features that it actually revealed to us a completely new body plan that existed during the Mesozoic. But it's also strange because it is a member of a group considered for a long time well-known and over-studied – the dromaeosaurids— although the recent discovery of “aberrant," unusual South American dromaeosaurids (the unenlangiines) already hinted at the fact that there is still much to discover and learn about these. Yet it is so different from any other known dromaeosaurid that it suggests we're still far from fully understanding the real diversity of the group.

佐：巴拉乌尔龙的异常与奇怪首先是由于其形态的多样性，有着太多的独一无二的特点，实际上它向我们展示了一种在中生代的全新身体结构。但它的奇怪也是由于它是一种被认为有了很长时间的了解和深入研究的物种——厉爪属恐龙——的成员，虽然最近发现的一种“异常”的南美厉爪属恐龙（unenlangiines）已经提醒人们有关这些物种还有许多未知需要发现和研究的事实。不过它和其他已知的厉爪属恐龙如此截然不同，表明我们依然离完全理解这一种群的真正的多样性相去甚远。

Unlike other contemporaries from the Romanian paleo-islands, that were strange because they were like their relatives from much earlier times, but unlike those living in the same time (kind of “living fossils” of their times, such as the duckbill Telmatosaurus or the armored Struthiosaurus), Balaur is unlike BOTH its predecessors and its contemporaries. It is simply unique...for now, at least.

和罗马尼亚古代岛屿上的其他同代生物不同，巴拉乌尔龙奇怪在他们和更早时期的近亲更相似，却和生活在同一时代的生物不同（某种那个时代的“活化石”，就像鸭嘴兽和厚甲龙），它们和前辈与同代都不同。光是这点就很独特了……至少目前如此。

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