原题：NOTES AND OBSERVATIONS ON COURTSHIP AND MATING IN Tityus (Atreus) magnimanus POCOCK, 1897 (Scorpiones: Buthidae) 

作者：Ross LK

本文详细记录了硕螯戾蝎交配前后及过程中的各个细节，对于了解硕螯戾蝎交配中的不同动作含义有重要的参考意义！

ABSTRACT: Courtship and mating behaviors of the scorpion Tityus (Atreus) magnimanus are herein described, consisting of various components that pertain to four distinct behavioral stages. The courtship and mating rituals of Tityus (Atreus) magnimanus are similar to those of other scorpions. Behavioral components are presented in an ethogram to demonstrate their occurrence during mating sequences. The current report is presented as observational data that were acquired during life history studies of this species.

摘要：本文描述了硕螯戾蝎的求偶和交配行为，包括了处于四个不同行为阶段的各种组成部分。硕螯戾蝎的求偶和交配仪式与其他蝎子相似。行为成分呈现在一个图谱中，以展现它们在交配序列中的出现。目前的报告是在该物种的生命历程研究中获得的观测数据。

INTRODUCTION

Courtship and mating among scorpion species, on which such data are available, comprise a series of complex, ritualized and characteristic behaviors that involve a courtship period of stereotyped acts leading up to sperm transfer from male to female via an external stationary spermatophore (1-5). Courtship in scorpions is usually divided into four distinct behavioral stages (different terminologies have been applied to individual stages by various authors): initiation, promenade á deux, sperm transfer and separation (3, 4, 6).

简介

现有数据表明，蝎子的求偶和交配包括一系列复杂的、仪式化的独特行为，这些行为包括求偶期的千篇一律的行为，这些行为导致精子通过外部固定的精荚从雄性转移到雌性。蝎子的求偶通常分为四个不同的行为阶段（不同作者对各个阶段使用了不同的术语）：起始、漫步（注：该词经查为葡萄牙语“双人漫步”，即公拉母游荡的阶段）、精子转移和分离。

In the majority of scorpion species, courtship is initiated by males and typically involves a male approaching a female and grasping her chelae within his own (in some instances, a chelicerae-to-chelicerae grasp may be initiated by males and used simultaneously with the chelal grasp to control female movements). After the chelal grasp, the male initiates and guides the female in a series of coordinated movements termed promenade á deux. This courtship “dance” may last from five minutes to several days, with most courtship and mating sequences lasting from 30 to 60 minutes (3).

对大多数种类的蝎子而言是由雄性发起求爱行为，通常包括雄性接近雌性并抓住她的钳子（在某些情况下，钳子对钳子的抓握可能是由雄性发起的，同时“钳握”（注：应为须肢紧握，为便于理解可译为钳握）用于控制雌性的动作）。在钳握之后，雄性发起并引导雌性进行一系列被称为promenade á deux（双人漫步）的协调动作。这种求偶“舞蹈”可能持续5分钟到几天，（注：一直以为持续几天的情况是交配失败了于是强制把公蝎拿出来打断交配……图样图森破）大多数求偶和交配过程持续30到60分钟。

During the promenade, the male seeks out a suitable surface or structure upon which to deposit his spermatophore. Once the spermatophore has been extruded and attached to a suitable surface, the male then directs the female movements until her genital opening is positioned above the spermatophore. A brief struggle ensues and the female rocking motion assists in engaging the spermatophore with the genital operculum, and the sperm is released into this opening (7). Once sperm transfer is complete, the male usually initiates separation of the participants (3).

在“双人漫步”的过程中，雄性会寻找一个合适的表面或结构来放置精荚。一旦精荚被挤出并附着在合适的表面上，雄性就会引导雌性的运动，直到她的生殖器开口位于精子的上方。随后是短暂的挣扎，雌性的摇摆动作帮助精荚与生殖器盖接触，精子被释放到这个开口。一旦精子转移完成，雄性通常会和雌性分开。

Courtship and mating behaviors have been described in approximately thirty scorpion species (3, 4), representing seven of the 18 extant families (8). Based on data reviewed by Polis and Sissom (3), both rituals have been described in more species (n = 12) from the family Buthidae than for any other scorpion family. Tityus C. L. Koch, 1836 is the largest genus within this family, represented by 181 species (9, 10). Tityus and the genus Centruroides Marx, 1890 (Buthidae), represent the only scorpion genera recognized as being medically significant in the Nearctic and Neotropical regions (11-13).

大约有30种蝎子被描述了求偶和交配行为，它们代表了现存18个科中的7个科。根据Polis和Sissom的数据，这两种仪式在Buthidae科（杀牛蝎科/钳蝎科）的物种中被描述的比在任何其他蝎子科中都多(n = 12)。戾蝎属是该科最大的属，共有181种（注：截止目前至少已有240种）。戾蝎属和似刺尾蝎属被认为是在新北界和新热带界仅有的具有医学意义的蝎子属。

Despite the large number of species among Tityus, its abundance in many urbanized and rural human occupied areas, and the medical significance of many of its component species, courtship and mating in members of this genus are few in number (1, 14, 15).

尽管戾蝎属的物种数量众多，在许多城市化和农村人类居住的地区都很丰富，而且其许多组成物种具有医学意义，但该属成员的求爱和交配数量很少。

The following contribution is based on laboratory observations of 12 mating sequences in the Venezuelan endemic scorpion, Tityus (Atreus) magnimanus Pocock, 1897 (Buthidae), conducted during February and March 2008. Tityus (Atreus) magnimanus inhabits tropical altitudinal forests in the states of Falcón and Lara in north-central Venezuela, and is considered to be of medical importance, with all severe and near-fatal envenomations in Falcón state attributed to this species. The present work is part of a continuing study on the life history of Tityus (Atreus) magnimanus and represents the first report on this medically significant species.

以下投稿基于2008年2-3月期间对委内瑞拉特有的硕螯戾蝎的12个交配流程的实验室观察。硕螯戾蝎生活在委内瑞拉中北部Falcón州和Lara州的热带高纬度森林中，被认为具有重要的医学意义，Falcón州所有严重和近乎致命的中毒都归因于该物种。目前的工作是对硕螯戾蝎生命周期的持续研究的一部分，是对这一医学上重要物种的首次报道。

MATERIALS AND METHODS

In order to acquire female specimens of known age for life history studies, specimens of Tityus (Atreus) magnimanus were mated in the laboratory and observational data were recorded, where specimens of Tityus (Atreus) magnimanus (n = 84) were reared to maturity. Upon attaining sexual maturity, unrelated adult males (n = 12) and laboratory reared females (n = 12) were randomly selected and transferred to individual containers.

材料与方法

为了获得已知年龄的雌性标本用于生命周期研究，在实验室对硕螯戾蝎样本进行了交配并记录了观察数据，饲养了84只硕螯戾蝎样本至成熟。在性成熟后，随机选择不相关的成年雄性(n = 12)和实验室饲养的雌性(n = 12)，并将其转移到单独的容器中。

Mature females were housed in circular 9 x 23 cm clear plastic containers, provided with a 50 mm layer of moist potting soil, a piece of cork or shelter and several flat rocks. As the containers of females would also serve as mating arenas, the addition of flat rocks provided a surface structure for spermatophore deposition by males.

性成熟的雌性被安置在9x23厘米的圆形透明塑料容器中，提供50毫米厚的潮湿盆栽土层、一块软木或其他遮蔽物、和几块平坦的岩石。由于雌性的容器也可以作为交配场所，所以增加的平坦岩石为雄性的精荚粘附提供了一个表面。

Adult males were kept in individual 15 x 15 x 15 cm clear plastic containers, with a 50 mm layer of moist potting soil and a single piece of cork for shelter. All specimens were maintained at temperatures from 26.6 to 29.0°C during the day and 23.8 to 25°C at night. The substrate was kept moist while ambient humidity levels were maintained in the range of 70 to 80%. Prey consisted of adult house crickets (Acheta domesticus L.) and cockroaches (Blatta lateralis Walker) with the type of prey alternated between ad libitum feedings. All observations were conducted under a low-intensity 60 watt red incandescent bulb in a parabolic reflector, suspended 50 cm above breeding arenas.

成年雄性被单独饲养在15x15x15厘米的透明塑料容器中，容器中有50毫米的湿润盆栽土壤和一块软木作为遮蔽物。所有样本白天保持在26.6-29.0℃，夜间保持在23.8-25℃。环境湿度保持在70-80%的范围内，基底保持湿润。食物以成体蟋蟀和樱桃蟑螂为主，两种食物随意交替饲喂（注：ad libitum德语：随意地）。所有的观察都是在一个低强度的60瓦红色白炽灯泡的抛物反射面下进行的，该反射器悬挂在繁殖场所上方50厘米处。（注：蝎子对红光不敏感，相关的资料正在学习如GREGORY R. C.等人关于不同波长光线对蝎子的活动影响（doi:10.1016/j.anbehav.2007.12.022）。也有Arachnoboard论坛上的网友表示蝎子根本看不见红光，但我暂时没找到相关文献。）

Mature virgin females were provided a 24-hour acclimation period before mating trials were initiated. A randomly selected mature male was introduced into each of 12 breeding arenas and all observed courtship and mating behaviors were recorded.

在开始交配试验之前，给予性成熟的处女雌性24小时的环境适应期。在12个繁殖场所中，随机选择一只成年雄性，记录所有观察到的求偶和交配行为。（注：这只公可真是不虚蝎生啊）

Once sperm transfer was completed, indicating termination of the mating sequence, males were removed from arenas and returned to individual containers.

一旦精子转移完毕，这意味着交配过程结束了，公蝎从交配场所中被拿回单独的容器里面。

RESULTS

In each of the 12 observed mating sequences, males immediately initiated courtship upon contact with females. As has been previously reported in scorpions (3), four distinct behavioral stages were observed: initiation, promenade á deux, sperm transfer, and separation. If specimens became separated during the promenade á deux, the behavioral sequence was immediately resumed by specimens without a return to a previous stage. The following ethogram demonstrates each behavioral component of courtship and the order in which they occurred during mating sequences (Figure 1).

结果

在12组交配进程中，每组的公蝎见到母蝎后马上就开始求爱，正如之前在参考文献3中报道的一样，观察到四个不同的行为阶段：起始、双人漫步、精子转移和分离。如果样本在双人漫步期间被分离，则样本立即恢复继续中断的行为，而不会返回到前一阶段。下表展示了求偶过程中每个行为组成部分以及它们在交配序列中发生的顺序（表1）。

Stage I: Initiation

In all observed mating sequences, males initiated courtship after detecting the movements of females (n = 3; 25%) or during random encounters with stationary females (n = 9; 75%). When a male detected a moving female or encountered a stationary female, courtship was initiated by the male grasping and rapidly moving his chelae over the segments (including the chelae) of one (n = 8; 66.6%) or both (n= 2; 16.7%) pedipalps of a female.

第一阶段:起始

在所有观察到的交配序列中，雄性在发现雌性的运动后开始求爱(n=3；25%)或随机遇到静止的雌性(n=9；75%)。当雄性发现一只移动的雌性或遇到一只静止的雌性时，雄性就会抓住并迅速移动它其中一个钳子(n=8；66.6%)或两个（n=2；16.7%)。

Two males (16.7%) encountered immobile females and immediately grasped and moved their chelae over metasomal segments III to V of each female. After a brief period (less than 45 seconds) of metasomal or pedipalpal grasping behaviors, a male would grasp each chela of a female within his own and align his body with that of the female in a chelicerae-to-chelicerae position thus, making contact with the female and terminating the initiation stage of courtship.

2只雄性(16.7%)遇到静止的雌性，立即抓住并将它们的钳子移动到每一只雌性的第III至V节后体上方。在短暂的一段时间内(少于45秒)的钳住钳子或后体的行为之后，雄性会抓住雌性的双钳，并将自己的身体与雌性以“钳子对钳子”的方式对齐，从而与雌性接触，结束求爱的起始阶段。

After initial contact by males, females typically withdrew their pedipalps against their own bodies and remained immobile until males initiated pulling movements. If a female resisted the attempts of a male to begin directed movements during initial contact, the male would judder for 10 to 25 seconds, then vigorously and repeatedly pull the female toward him until she began moving.

在与雄性进行初次接触后，雌性通常会将须肢缩回到自己的身体上并保持不动，直到雄性开始拉扯动作。如果雌性在初次接触时拒绝雄性的引导动作，雄性会颤抖10到25秒，然后用力反复地将雌性拉向自己，直到雌性开始移动。

Polis and Sissom (3) reported that juddering by male scorpions functions either as a highly ritualized sexual or species-recognition behavior, as an instigator of mating behavior so that the female is stimulated to cooperate, or as a simple byproduct of intense sexual excitation. The only instances in which juddering was exhibited by males was in direct response to resistant females during the initiation and promenade á deux stages and appeared to be used by males to suppress unreceptive behaviors and stimulate females into mating.

Polis和Sissom报告说，雄蝎子的抖动，要么是一种高度仪仪化的性行为或是是一种物种识别行为——作为交配行为的煽动者，以此刺激雌性进行协作交配，要么只是强烈性兴奋的简单表现。雄虫表现出抖动的唯一情况是在起始期和“双人漫步”期对雌性抵抗的直接反应，似乎是雄虫用来阻止雌性不接受的行为和刺激雌性交配。

In two pairings that involved receptive females that never resisted males during courtship movements, juddering was not observed in either male throughout all stages of courtship. These observations suggest that juddering behavior may be used by courting males to stimulate unreceptive females into mating.

在两组求偶过程中从不抗拒雄性的雌性交配中，在求偶的所有阶段都没有观察到雄性的抖动。这些观察结果表明，抖动行为可能是求爱的雄性用来刺激不接受交配的雌性。

Alexander (1) reported metasomal grasping by courting males of Tityus (Atreus) trinitatis Pocock, 1897 and suggested that such behaviors by males may function as a means of identifying conspecific females through the sexually differentiated morphologies of metasomal segments. Repeated grasping of pedipalps and chelae of females by courting males may serve as a similar means of identifying conspecific females via differences in these morphological structures.

Alexander报道了求偶雄性硕螯戾蝎的“后体抓握”行为，并认为雄性的这种行为可能是通过后体的性别分化形态来识别同种雌性的一种手段。雄性在向雌性求爱时反复抓住须肢和钳子，可能是通过这些形态结构的差异来识别同种雌性的一种类似手段。（注：通过网上各种信息的搜寻，国内的蝎子饲养者普遍认为蝎子的“控尾”——即文中的“后体抓握”行为是雄蝎让雌蝎冷静、保护自己的手段。个人比较赞同该观点。因为仔细观察，雌雄碰面甚至仅仅同处一缸感知到对方的存在之时，双方有一个“发呆期”，此间双方都在用栉状器拍打摩擦地面，甚至抖动身体来发送信号，此时应该已经完成了对对方物种的判断，没有必要再以生命为代价去识别后体。）

Sexually mature males and females of Tityus (Atreus) magnimanus exhibit differing morphologies of the pectines, chelae and metasomal segments. As has been reported by Lourenço (9), females of some species within the subgenus Atreus Gervais, 1843 possess enlarged basal middle lamellae that are not present in males. The chelal manus of males is moderately inflated and serves to differentiate mature males from immature males and females. Additionally, females are larger and more robust than males and the metasomal segments of sexually mature females are thickened and well developed in comparison to those of males.

性成熟的雌雄硕螯戾蝎的栉状器、钳子和后体都表现出不同的形态。正如Lourenço所报道的那样，1843年，一些Atreus亚属（注：现为Scorpio，蝎属）的雌性具有扩大的骨板中间片（注：即栉状器根部有“大齿”），而雄性没有。雄性的钳子有适度的膨胀，用以区分成熟雄性与不成熟雄性、雌性。此外，雌性比雄性更大、更健壮，性成熟雌性的后体比雄性更厚、更发达。

 

Stage II: Promenade á Deux

The promenade á deux occurs in all scorpion species for which courtship and mating data are available (2, 3, 4). As in other scorpion species, the males of Tityus (Atreus) magnimanus direct and coordinate the movements of the pair during the promenade á deux. In Tityus (Atreus) magnimanus, the promenade begins when a male firmly grasps the chelae of a female within his own and begins to guide the female in mutually backward, sideways and rarely, forward movements. The male begins promenade movements by elevating its body slightly above the ground surface with the metasoma elevated and dorsomedially flexed, and moving backwards, effectively pulling the female along and directing her movements.

第二阶段:双人漫步

在所有有求偶、交配数据的蝎子物种中，双人漫步都会发生。与其他蝎子物种一样，雄性硕螯戾蝎在双人漫步期间指导和协调配偶的行动。对硕螯戾蝎而言，当一只雄性牢牢抓住一只雌性的螯，并开始引导雌性相互向后、向侧面移动，很少向前移动时，漫步就开始了。雄性开始散步时，身体略高于地面，后体抬高并向背部弯曲，然后向后移动，有效地拉着雌性前进，指导她的动作。

During backwards movements, the male will periodically flex and straighten the metasoma while simultaneously raising and lowering it in response to the willingness of the female to follow his movements.

在向后运动的过程中，雄性会周期性地弯曲和伸直后体，同时，为了让雌性跟随自己的运动，他们会根据雌性的意愿同步地抬起和放下后体。

If a female resists or stops moving, the male will move closer to her, elevate his body, and begin pulling the female in vigorous backward movements while simultaneously elevating and waving his metasoma from side-to-side. Females typically keep their bodies close to the substratum with the metasoma flexed and held laterally.

如果雌性抵抗或停止移动，雄性会靠近她，抬起身体，开始以有力的向后动作拉着雌性，同时抬起并左右摆动他的后体。雌性通常保持它们的身体靠近基底，而后体弯曲并保持在侧面。

During this stage of courtship, the male guides the female across the substratum until a suitable surface is located for spermatophore deposition. During promenade movements, male pectines are directed downward and are moved back and forth over the substratum and all encountered surface structures. During promenade movements, the female may resist male attempts to direct coordinated movements of the pair by pulling and trying to move in a direction opposite that of the male or by ceasing movements, lowering her body to the substratum and becoming still.

在求爱的这一阶段，雄性会引导雌性走过基底，直到找到一个适合存放精荚的表面。在双人漫步中，雄性向下在基底和所有遇到的表面结构上来回移动。在散步运动中，雌性可能会抵制雄性试图引导其协调运动的尝试，通过拉拽并试图朝着与雄性相反的方向移动，或者停止运动，将身体降至基底并保持静止。

Immobile females only resume moving after males exhibit juddering (6 to10 seconds) and vigorous pulling behaviors. In all observed pairings, the promenade á deux lasted from 4.5 to 9.0 minutes and was terminated when a suitable location for spermatophore deposition was selected by the male. A courting pair may pass over a structure several times from different directions until the male ceases their movements. Selection of a proper structure for spermatophore deposition finishes the promenade á deux.

静止不动的雌性只有在雄性表现出抖动（6~10秒）和有力的拉扯行为后才能恢复活动。在所有观察到的交配中，双人漫步持续时间为4.5~9.0分钟，当雄性选择合适的精荚放置位置时终止。一对求偶的蝎子可能会从不同的方向多次经过一个基底，直到雄性停止它们的动作，为放置精荚选择一个合适的结构，双人漫步才完成。

 

Stage III: Sperm Transfer

As in other scorpions, sperm transfer in Tityus (Atreus) magnimanus is accomplished via a stationary spermatophore that is flagelliform in shape and structure (7). Males of this species are capable of producing their first spermatophore within seven days of maturing molts and can regenerate spermatophores in a very short period, 24 hours (Ross, personal observation).

与其他蝎子一样，硕螯戾蝎的精子转移是通过一个固定的鞭状精荚完成的。雄性硕螯戾蝎能够在蜕皮成熟后的7天内产生第一个精荚，并能在很短的时间内再生精荚，24小时(来自Rose的个人观察)。

Once a suitable surface is selected by a male, he pulls the female close and with his chelicerae grasps either the anterior edge of her carapace (n = 10; 83.3%) or chelicerae (n = 2; 16.7%). The courting pair remains still during the release of the spermatophore. The male lowers his venter against the selected surface and the pedicel is attached to the surface. Whereas the spermatophore is slowly extruded, the male gradually elevates his body while simultaneously moving backwards until the trunk and flagellum of the spermatophore are exposed.

一旦雄性选择了一个合适的表面，它就会把雌性拉近，用它的钳子抓住它的甲壳的前缘（n = 10;83.3%）或螯肢（n = 2;16.7%）（注：很纳闷是怎么抓住的，雌蝎把牙伸出来让雄蝎抓吗？……）。在精子被释放的过程中，这对求偶的蝎子保持静止不动。雄蝎的腹部向选定的表面降低让精荚附着在表面上。当精荚被缓慢挤出时，雄性逐渐抬起身体，同时向后移动，直到精荚的茎部和鞭毛露出来。

Spermatophore expulsion is marked by the male elevating his entire metasoma vertically and slowly waving it from side-to-side (15 to 39 seconds). Once the spermatophore is deposited, male metasomal movements cease (n = 12; 100%) and he pulls the female toward him and directs her movements until her genital aperture is positioned directly above the erect spermatophore. Once the female genital orifice is positioned above the capsule, the male will rock the female back and forth (6 to 10 seconds) so that the spermatophore is engaged by her genital operculum. During the back-and-forth rocking movements, the female lowers her body upon the trunk of the spermatophore that houses the sperm vesicle and sperm duct, so that the sperm mass is released into her genital aperture. In this observation, as soon as the sperm mass was transferred, females (n = 12; 100%) immediately separated and retreated 4 to 10 cm away from males.

精荚排出的标志是雄性将整个后体垂直抬起，慢慢地从一边到另一边摆动（15~39秒）。一旦精荚被放置，雄性后体运动停止（n =12;100%），雄性将雌性拉向自己并引导她的动作，直到她的生殖器孔位于竖直的精荚的正上方。一旦雌虫的生殖孔位于荚膜上方，雄虫就会前后摇动雌虫（6~10秒），这样精荚就会被雌蝎的生殖盖包裹住。在前后摇摆的过程中，雌性把身体放在装有精囊和精管的精荚茎部上，这样精子团就被释放到她的生殖孔中。在这项观察中，精子一经转移，雌性（n =12;100%）立即分开，并退到离雄性4~10厘米远的地方。

 

Stage IV: Separation

In all sequences in the present study, mating terminated when the female pulled away and moved a short distance from the male immediately following sperm transfer. As soon as a female started to struggle and pull away, a male simply released his grasp upon female. Once a female moved away from a male, she either rubbed her venter upon the substratum (n = 7; 58.3%) or used one or both pairs of anterior walking legs to vigorously rub her genital operculum (n = 5; 41.7%).

第四阶段:分离

 

在本研究的所有序列中，当雌性在精子转移后立即抽离并离开雄性一小段距离时，交配就终止了。一旦雌性开始挣扎并挣脱，雄性就会松开对雌性的控制。一旦雌性离开雄性，她要么在基底上摩擦她的生殖器（n=7；58.3%）或用一只或一对前腿用力摩擦生殖盖（n = 5；41.7%）。

The male remained with a spent spermatophore and rapidly moved the tarsi of the first pair of walking legs over the base and trunk of the spermatophore (n = 9: 75%) (4 to 8 seconds) or remained stationary without exhibiting any activity (n = 3; 25%) (30 to 120 seconds).

雄蝎留下一个用过的精荚，在精荚的根部和躯干上快速移动第一对行走腿的跗足（n=9.75%）（4 ~ 8秒），或者静止不动（n = 3;25%）（30 ~ 120秒）。

After the recuperative stage, males (n = 12; 100%) moved slightly backwards and used their chelicerae to sever the flagellum near its base on the spermatophore. After severing the flagellum, males rested for a brief period (2 to 6 minutes) before becoming vagile and moving away from spermatophores. Consumption of the spermatophore by male or female was not observed in any mating sequence. Post- mating males and females do not exhibit agonistic or aggressive behaviors toward each other and will aggregate in large, mixed gender groups in laboratory enclosures.

休息期过后，雄性（n=12;100%）稍微向后移动，用它们的螯肢切断位于精荚基部附近的鞭毛。在剪去鞭毛后，雄性会休息一段时间（2至6分钟），然后开始漫游并离开精荚。在任何交配顺序中都没有观察到雄性或雌性对精荚的消耗。（注：一直以为是吃精荚，见发布的视频）交配后的雄性和雌性不会对彼此表现出敌对或攻击行为，并且会聚集在实验容器中的大型混合性别群体中。

 

DISCUSSION

The courtship and mating behaviors of Tityus (Atreus) magnimanus are similar to behaviors described in other scorpions including several species within the genus Tityus. However, in order to explain and more thoroughly understand patterns of phylogeny, ecology, biology, reproductive evolution and diversification among scorpions, a precise knowledge of the reproductive biology and associated behaviors in scorpions is required (12).

讨论

 

硕螯戾蝎的求偶和交配行为与其他蝎子（包括戾蝎属的几个物种）的行为相似。然而，为了解释和更彻底理解蝎子的系统发育、生态学、生物学、生殖进化和多样化模式，需要对蝎子的生殖生物学和相关行为有精确的了解。

The lengthy and time consuming process of the promenade á deux has been traditionally interpreted as the time necessary for a courting male to locate and select a suitable surface upon which he will deposit his spermatophore. However, when a courting male engaged in the promenade á deux is presented with two different surface structures (a stick and a flat rock) during successive mating trials, the male will select the flat rock instead of the stick for spermatophore deposition during one trial and when introduced into another mating arena with a new female, chooses the stick instead of the rock.

这一漫长而耗时的过程传统上被解释为求偶的雄性寻找并选择一个合适的表面来放置他的精荚所需要的时间。然而，在连续的交配试验中，对在双人漫步中的求偶雄性，在连续的交配试验中呈现两种不同的表面结构（枯枝和平坦的岩石）时，它们会在一次试验中选择平坦的岩石而不是枯枝来放置精荚，当被引入另一个与新雌性交配的场所时，选择枯枝而不是岩石。

Additionally, a male may direct the female over a surface dozens of times before finally selecting the surface as the site of spermatophore deposition. Francke (7) reported that males of Centruroides sculpturatus Ewing, 1928 (Buthidae) when maintained overnight in plastic bags with pieces of crumpled newsprint for cover would deposit spermatophores upon the newsprint.

此外，雄性在最终选择一个表面作为精荚放置的地点之前，可能会引导雌性在一个表面上经过几十次。Francke曾报道，雄性雕纹似刺尾蝎放在塑料袋中过夜时，里面放一块皱报纸盖住，它会将精荚放置在报纸上。

Overall, a courting male does not appear to be overly selective in his choice of site for spermatophore deposition. Personal observations of additional mating sequences in Tityus (Atreus) magnimanus and Tityus (Atreus) trinitatis Pocock, 1897 have revealed that a courting male will select any solid surface encountered for spermatophore deposition including compacted substrate, fragments of clay plant pots, pieces of peat seedling pots, rocks, sticks, various tree barks, dried leafs, sections of palm root, quarter dollars, plastic bottle caps, cardboard and construction paper.

总的来说，求爱的雄性在选择精荚放置的地点时似乎没有过度的选择性。通过个人对硕螯戾蝎和特立尼达戾蝎的额外交配序列观察发现，求偶的雄性会选择任何遇到的固体表面来放置精荚，包括压实的基质、粘土花盆的碎片、泥炭苗盆的碎片、岩石、树枝、各种树皮、干叶子、棕榈根的部分、25美分硬币、塑料瓶盖、纸板和建筑纸。

Male juddering has been reported to occur in 18 species, from eight families, and has been reported to occur during all stages of courtship (3, 4). Juddering behavior during the promenade á deux was observed in ten pairings of Tityus (Atreus) magnimanus and was utilized by a male in response to a female that became unresponsive to male-directed movements or attempted to move in an opposite direction of male during courtship. In two pairings, females did not resist male- directed movements or attempt to pull away and move in an opposite direction. In both pairings, juddering was not displayed by either male during any stage of courtship. These observations suggest that juddering by male Tityus (Atreus) magnimanus may be used in direct response to resistance behaviors exhibited by females during the promenade and may act to stimulate mating.

据报道，雄性抖动发生在8个科的18个物种中，并且发生在求偶的所有阶段。在10对硕螯戾蝎中观察到在双人漫步期间的抖动行为，这是雄性对雌性在求偶期间对雄性引导的动作没有反应或试图向相反方向移动的反应。在两对交配蝎子中，雌性没有抵抗雄性的动作，也没有试图抽离并朝相反的方向移动。这两对中的雄性在求偶的任何阶段都没有表现出抖动。这些观察结果表明，雄性硕螯戾蝎的抖动可能是对雌性在散步时表现出的抵抗行为的直接反应，也可能是刺激交配的行为。

Juddering by the male during the initial stage did not occur in any of the 12 pairings observed during this study. Males introduced into mating arenas containing females either sensed their movements and moved directly to them or encountered immobile females during random movements. Males did not exhibit pre-contact mate recognition behaviors and readily approached moving and immobile females. Once contact was made with a female, the male would immediately grasp a single pedipalp of the female and make repeated grasps upon the various segments. In two pairings, the male made repeated grasps upon metasomal segments III to V of the female.

在这项研究中观察到的12对配对中，雄性在初始阶段没有出现抖动。被引入有雌性的交配场所的雄性要么感觉到它们的动作并直接向它们移动，要么在随机移动中遇到不动的雌性。雄性没有表现出接触前的配偶识别行为，并且很容易接近活动和不活动的雌性。一旦与雌性接触，雄性会立即抓住雌性的一条须肢，并反复抓住不同的部分。在两对配对中，雄性反复抓住雌性的第III到V节后体。

Alexander (1) noted that there is a marked sexual dimorphism of the metasomal segments in males and females of Tityus (Atreus) trinitatis and that courting males would approach females and repeatedly grasp various metasomal segments. Furthermore, the author suggested that males utilize grasping behaviors to identify the sexually dimorphic characters of the metasomal segments in females. The exhibition of grasping behaviors in courting males of Tityus (Atreus) magnimanus and Tityus (Atreus) trinitatis propose that differences in the morphology of various structures (chelae, pedipalps, metasomal segments) are important for mate recognition by males of both species during courtship.

Alexander注意到特立尼达戾蝎的雄性和雌性之间存在着明显的两性二态性，求偶的雄性会接近雌性并反复抓住后体的不同节。此外，他表示，雄性利用抓握行为来识别雌性后体关节的两性二态特征。硕螯戾蝎和特立尼达戾蝎在求偶过程中表现出的抓握行为表明，两种雄性在求偶过程中各种结构（螯、须肢、后体关节）的形态差异对它们的择偶识别很重要。