Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea
Исследования относительного роста частей тела морской свиньи впервые показали, что он в постнатальном онтогенезе существенно различается в четырех секторах, соответствующих черепу и отделам позвоночника. Наибольшими темпами относительного роста отличаются грудной и хвостовой отделы. Средняя часть те...
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Інститут зоології ім. І. І. Шмальгаузена НАН України
2005
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| Cite this: | Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea / P. E. Gol’din // Вестн. зоологии. — 2005. — Т. 39, № 5. — С. 59–65. — англ. |
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| author | Gol'din, P. E. |
| author_facet | Gol'din, P. E. |
| citation_txt | Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea / P. E. Gol’din // Вестн. зоологии. — 2005. — Т. 39, № 5. — С. 59–65. — англ. |
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| description | Исследования относительного роста частей тела морской свиньи впервые показали, что он в постнатальном онтогенезе существенно различается в четырех секторах, соответствующих черепу и отделам позвоночника. Наибольшими темпами относительного роста отличаются грудной и хвостовой отделы. Средняя часть тела отличается наибольшими темпами относительного роста у обоих полов. Выдвинута гипотеза о том, что двухстадийный рост организма определяется периодичностью роста хвостового отдела. Половой диморфизм размеров тела создается за счет частей тела, проявляющих положительную аллометрию или изометрию у обоих полов, однако различия в абсолютных величинах затрагивают лишь анально-генитальное расстояние.
The studies of the allometric growth of the body parts in harbour porpoises have shown that the growth of body parts during the postnatal ontogenesis of the harbour porpoise differs substantially in four body sectors, which correspond to the skull and the parts of vertebral column. The highest rates of allometric growth are observed in the thoracic and caudal departments. The middle part of the body is characterized by maximum rates of allometry in both sexes. The hypothisis was brought forward that the two-stage growth of the organism is determined by the periodicity of growth of the caudal department. Sexual dimorphism of the body size is determined by the body parts, which exhibit positive allometry or isometry in both part. Sexual differences in growth rates are observed in the lumbar part but the differences of absolute length values concern only anusgenital distance.
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UDC 591.4:599.536(262.5)
BODY PROPORTIONS OF HARBOUR PORPOISE
PHOCOENA PHOCOENA (CETACEA, PHOCOENIDAE)
IN THE SEA OF AZOV AND THE BLACK SEA
P. E. Gol’din
V. I. Vernadsky Taurida National University,
Vernadsky Avenue, 4, Simferopol, AR Crimea, 95007 Ukraine
E-mail: oblako@home.eris.net
Accepted 31 November 2003
Ïðîïîðöèè òåëà ìîðñêîé ñâèíüè Phocoena phocoena (Cetacea, Phocoenidae) â Àçîâñêîì è ×åðíîì
ìîðÿõ. Ãîëüäèí Ï. Å. — Èññëåäîâàíèÿ îòíîñèòåëüíîãî ðîñòà ÷àñòåé òåëà ìîðñêîé ñâèíüè âïåð-
âûå ïîêàçàëè, ÷òî îí â ïîñòíàòàëüíîì îíòîãåíåçå ñóùåñòâåííî ðàçëè÷àåòñÿ â ÷åòûðåõ ñåêòîðàõ,
ñîîòâåòñòâóþùèõ ÷åðåïó è îòäåëàì ïîçâîíî÷íèêà. Íàèáîëüøèìè òåìïàìè îòíîñèòåëüíîãî
ðîñòà îòëè÷àþòñÿ ãðóäíîé è õâîñòîâîé îòäåëû. Ñðåäíÿÿ ÷àñòü òåëà îòëè÷àåòñÿ íàèáîëüøèìè
òåìïàìè îòíîñèòåëüíîãî ðîñòà ó îáîèõ ïîëîâ. Âûäâèíóòà ãèïîòåçà î òîì, ÷òî äâóõñòàäèéíûé
ðîñò îðãàíèçìà îïðåäåëÿåòñÿ ïåðèîäè÷íîñòüþ ðîñòà õâîñòîâîãî îòäåëà. Ïîëîâîé äèìîðôèçì
ðàçìåðîâ òåëà ñîçäàåòñÿ çà ñ÷åò ÷àñòåé òåëà, ïðîÿâëÿþùèõ ïîëîæèòåëüíóþ àëëîìåòðèþ èëè
èçîìåòðèþ ó îáîèõ ïîëîâ, îäíàêî ðàçëè÷èÿ â àáñîëþòíûõ âåëè÷èíàõ çàòðàãèâàþò ëèøü
àíàëüíî-ãåíèòàëüíîå ðàññòîÿíèå.
Êëþ÷å âûå ñ ë î â à: ìîðñêàÿ ñâèíüÿ, Àçîâñêîå ìîðå, ×åðíîå ìîðå, ïðîïîðöèè òåëà,
àëëîìåòðèÿ.
Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov
and the Black Sea. Gol’din P. E. — The studies of the allometric growth of the body parts in harbour
porpoises have shown that the growth of body parts during the postnatal ontogenesis of the harbour
porpoise differs substantially in four body sectors, which correspond to the skull and the parts of
vertebral column. The highest rates of allometric growth are observed in the thoracic and caudal
departments. The middle part of the body is characterized by maximum rates of allometry in both sexes.
The hypothisis was brought forward that the two-stage growth of the organism is determined by the
periodicity of growth of the caudal department. Sexual dimorphism of the body size is determined by
the body parts, which exhibit positive allometry or isometry in both part. Sexual differences in growth
rates are observed in the lumbar part but the differences of absolute length values concern only anus-
genital distance.
Ke y wo r d s: harbour porpoise, the Sea of Azov, the Black Sea, body proportions, allometry.
Introduction
Body proportions and regularities of allometric growth in the postnatal ontogenesis of harbour porpoise
were studied by several authors (Zalkin, 1938; Van Utrecht, 1978; Stuart, Morejohn, 1980; Read, Tolley,
1997); some results of these studies demonstrated a number of contradictions. Zalkin (1938) was the only
one who dealt with porpoises from the Sea of Azov and the Black Sea; however, his research was impeded
by the lack of age determination methods. The objective of this study is to describe the patterns of allometry
in porpoises from the Black Sea basin and their changes due to postnatal growth.
Material and methods
The material for this study was obtained from 125 harbour porpoises: 97 from the Sea of Azov and 28
from the Black Sea, found dead at the coastline or by-caught during the fisheries operations in 1997–2003.
The data on certain measurements taken from 25 animals were kindly provided by V. V. Pavlov (S. I. Geor-
Vestnik zoologii, 39(5): 59–65, 2005
© P. E. Gol'din, 2005
Ìîðôîëîãèÿ
gievsky, Crimean State Medical University), data on 5 animals by N. V. Frolova (Donetsk National
University). The rest of the material was collected by the author based on Department of Zoology,
V. I. Vernadsky Taurida National University.
Morphometric measurements were taken according to complete or incomplete protocol (fig. 1). The
main measurements were taken twice, from the opposite points (tip of rostrum and notch of flukes), to reduce
the error of measuring.
Sex was determined in 117 specimens, age in 119 specimens. The age was determined by counting
growth layer groups (GLGs) in dentine according to the common techniques, with the use of thin longitu-
dinal sections of decalcified teeth stained by Erhlich’s or Mayer’s haematoxylin (see details in: Gol’din, in
press). The age of 3 specimens was determined from the GLGs number in mandible using the technique
developed by author (Gol’din, 2003).
The growth curves were calculated using Gompertz formula as follows:
,
where t is age in years, L is body length in cm, b and k are constants, L is asymptotic length in cm, as well
as models including several Gompertz formulae.
The allometry equations were calculated using power formula as follows:
y = axb.
The indices of morphometric measurements were calculated as their ratios to the total body length.
Results and discussion
The main morphometric measurements in harbour porpoise can be compared with
the positions of certain skeleton elements. The anterior edge of a flipper basis is locat-
ed approximately at the level of the first thoracic vertebrae. The anterior edge of the
dorsal fin is located at the level of one of the first lumbar vertebrae (from L1 to L5,
the most often at L1 or L4); the posterior edge is from L5 to L13, the most often at
8
60 P. E. Gol'din
Fig 1. Morphometric measurements of the harbour porpoise: L — total body length, 1 — rostrum to tip of
mouth; 2 — rostrum to eye; 3 — rostrum to blowhole; 4 — rostrum to axilla; 5 — rostrum to front edge of
base of dorsal fin; 6 — rostrum to back edge of base of dorsal fin; 7 — rostrum to centre of genital slit; 8 —
rostrum to centre of anus; 9 — centre of genital slit to centre of anus (anus-genital distance); 10 (Lpect) —
length of flipper (max); 11 — width of flipper (max); 12 — base of dorsal fin; 13 — height of dorsal fin;
14 — length of outer edge of tail fluke (from the tail axis); 15 — maximum width of tail fluke; 16 — maximum
fluke span; I— IV — number of Sector. Longitudinal measurements are indicated only from tip of rostrum.
Ðèñ. 1. Ñõåìà ìîðôîìåòðè÷åñêèõ èçìåðåíèé ìîðñêîé ñâèíüè: L — îáùàÿ äëèíà òåëà; 1 — îò ðîñòðó-
ìà äî óãëà ðòà; 2 — îò ðîñòðóìà äî ïåðåäíåãî óãëà ãëàçà; 3 — îò ðîñòðóìà äî ïåðåäíåãî êðàÿ äûõàëà,
4 — îò ðîñòðóìà äî ïåðåäíåãî êðàÿ ãðóäíîãî ïëàâíèêà; 5 — îò ðîñòðóìà äî ïåðåäíåãî êðàÿ ñïèííîãî
ïëàâíèêà; 6 — îò ðîñòðóìà äî çàäíåãî êðàÿ ñïèííîãî ïëàâíèêà; 7 — îò ðîñòðóìà äî öåíòðà ïîëîâîé
ùåëè; 8 — îò ðîñòðóìà äî öåíòðà àíàëüíîãî îòâåðñòèÿ; 9 — îò öåíòðà ïîëîâîé ùåëè äî öåíòðà àíàëü-
íîãî îòâåðñòèÿ (àíàëüíî-ãåíèòàëüíîå ðàññòîÿíèå); 10 (Lpect) — äëèíà ãðóäíîãî ïëàâíèêà (ìàêñè-
ìàëüíàÿ); 11 — øèðèíà ãðóäíîãî ïëàâíèêà (ìàêñèìàëüíàÿ); 12 — äëèíà îñíîâàíèÿ ñïèííîãî ïëàâ-
íèêà; 13 — âûñîòà ñïèííîãî ïëàâíèêà; 14 — äëèíà âíåøíåãî êðàÿ ëîïàñòè õâîñòà (îò õâîñòîâîãî
ñòåáëÿ); 15 — ìàêñèìàëüíàÿ øèðèíà ëîïàñòè õâîñòà; 16 — ìàêñèìàëüíûé ðàçìàõ ëîïàñòåé õâîñòà;
I–IV — íîìåðà ñåêòîðîâ. Ïðîäîëüíûå ïðîìåðû ïîêàçàíû òîëüêî îò êîíöà ðîñòðóìà.
L10 or L12. The anus is located approximately at the level of the front part of the cau-
dal department, Ca5–Ca8.
Thus, there can be distinguished four conventional sectors corresponding appro-
ximately to the skeleton departments:
I — between the tip of rostrum and the axilla; corresponds to the skull and the
cervical department of the vertebral column;
II — between the axilla and the basis of the dorsal fin; corresponds to the thoracic
department of the vertebral column;
III — between the basis of the dorsal fin and the anus; corresponds to the lumbar
department of the vertebral column;
IV — between the anus and the notch of the flukes; corresponds to the caudal
department of the vertebral column.
The lengths of Sectors III and IV are on an average 2 cm less than the correspon-
ding measurements of the vertebral column. This is particularly caused by the measur-
ing technique: the external measurements were taken point to point, and measurements
of the departments of the vertebral column were taken along the bends with the flexi-
ble measuring tape. The length of Sector II is on an average somewhat exceeds the
length of the thoracic department, because the former is often influenced by the back-
ward displacement of the dorsal fin in many specimens, and the bend in the thoracic
department is not clearly expressed.
Sector I, the anterior part of the body, is characterized by the negative allometry
(here and ff. see table 1). This tendency remains during all postnatal ontogenesis, since
the moment of birth. The index of Sector I (the ratio of the sector length to the body
length), which takes on an average 26.8 and 26.4% of the body length in neonate males
and females, decreases to 23.5 and 23.1% during the first year of life and reaches 22
and 21% in adults (here and ff. see table 2). Sector I takes only 19.2–21.1% of the body
length in the longest males and 18.1–20.6% in the longest females. It is interesting to
61Body proportions of harbour porpoise Phocoena phocoena
Ta b l e 1. Coefficients of allometry equation y = axb for body parts of the harbour porpoise from the Sea of Azov
Ò à á ëèö à 1. Êîýôôèöèåíòû óðàâíåíèÿ àëëîìåòðèè y = axb äëÿ îòäåëüíûõ ÷àñòåé òåëà ìîðñêîé ñâèíüè
èç Àçîâñêîãî ìîðÿ
Sector I (ïðîìåð 4) 38 1.0 0.69 0.06 0.77 33 1.34 0.63 0.06 0.78
Sector II 33 0.07 1.22 0.09 0.85 23 0.05 1.32 0.17 0.72
Sector III 19 0.17 1.07 0.18 0.68 16 0.27 0.73 0.21 0.47
Sector IV 20 0.28 1.02 0.09 0.88 20 0.19 1.10 0.08 0.95
Sector I+II (ïðîìåð 5) 35 0.60 0.94 0.04 0.94 30 0.60 0.94 0.07 0.86
Sector II+III 19 0.26 1.12 0.06 0.94 18 0.27 1.11 0.07 0.94
Sector III+IV 33 0.42 1.05 0.03 0.96 30 0.44 1.04 0.06 0.91
Measurement 1 17 0.72 0.56 0.16 0.44 19 0.74 0.55 0.10 0.63
Measurement 2 29 0.59 0.66 0.10 0.63 30 0.46 0.71 0.08 0.72
Measurement 3 17 2.39 0.36 0.13 0.34 20 0.52 0.69 0.11 0.68
Measurement 6 30 0.90 0.92 0.03 0.98 28 1.06 0.88 0.06 0.90
Measurement 7 17 2.24 0.69 0.09 0.80 16 0.71 0.98 0.04 0.98
Measurement 8 19 0.76 0.98 0.04 0.96 20 0.89 0.94 0.04 0.96
Measurement 9 13 1.6·10-8 4.35 0.48 0.88 16 0.25 1.07 0.39 0.35
Measurement 10 27 0.37 0.80 0.07 0.82 26 0.52 0.72 0.09 0.74
Measurement 11 20 0.27 0.66 0.14 0.54 19 0.21 0.72 0.08 0.81
Measurement 12 30 0.34 0.82 0.10 0.68 28 0.28 0.86 0.13 0.62
Measurement 13 14 0.35 1.12 0.25 0.63 11 0.37 0.66 0.23 0.48
Measurement 14 13 0.60 0.72 0.11 0.77 9 0.65 0.71 0.18 0.68
Measurement 15 24 0.25 0.77 0.18 0.43 20 0.32 0.71 0.13 0.61
Measurement 16 12 0.88 0.71 0.11 0.79 9 0.19 1.03 0.19 0.81
Measurement/body part
Males Females
n a b SEb r2 n a b SEb r2
note that in the examined Black Sea specimens the index of Sector I is remarkably less;
it takes on an average 20% of the body length in both sexes.
The data on the proportions of Sector I in the specimens from the Sea of Azov
match exactly the results obtained by V. I. Zalkin (1938).
The negative allometry expresses itself in the growth of all the measurements with-
in Sector I (tip of rostrum to edge of mouth, eye, blowhole). The coefficients of the
allometry equations for the distance from the tip of rostrum to the blowhole differ sig-
nificantly in males and females but there are no significant differences in the absolute
values of this measurement.
Sector II demonstrates positive allometry in both sexes. Its rapid growth continues
during the first year of life. The index of Sector II takes 20.7 and 19.1% in neonate
males and females, rises up to 23.4 and 24.0% by the end of the first year, and then
remains at the level about 23% in males. In females the index of Sector II grows up to
25–26% by the age of 2–3 years, and then decreases approximately to 24% due to
increase of the posterior part of the body. In adult animals Sector II demonstrates a
tendency to positive allometry in regard to the total body length; in some of the longest
males it reaches 27%, in females 29%. Sector II is one of the two sectors determining
the total body length.
As a result, the anterior part of the body constituted by Sectors I and II demon-
strates slightly negative allometry (b = 0,94). Zalkin (1938) considered this tendency
to be the main direction of the body proportions change in the ontogenesis. Van
Utrecht (1978), basing on the measurements made along the body contour, reported
the negative allometry pattern of the anterior part of the body; however, at the same
time he recorded isometry in males and positive allometry in females as for the mea-
surement from the blowhole to the basis of the dorsal fin. A. J. Read and K. A. Tolley
(1997) calculated the coefficients of the allometry equations for the measurements from
the tip of rostrum to the basis of the dorsal fin as 0.81 and 0.88 respectively and noted
the slightly negative allometry of the basis of dorsal fin. Galatius (personal communi-
cation) specially emphasizes the necessity to study separately the two zones in the ante-
rior part of the body, which correspond to Sectors I and II and exhibit dramatically
contrasting growth patterns.
62 P. E. Gol'din
Ta b l e 2. Indices of body parts in age groups of the harbour porpoise from the Sea of Azov calculated as per-
centages of total body length
Ò à á ëèö à 2. Èíäåêñû ðàçëè÷íûõ ÷àñòåé òåëà â âîçðàñòíûõ ãðóïïàõ ìîðñêîé ñâèíüè èç Àçîâñêîãî ìîðÿ,
âûðàæåííûå â äîëÿõ îáùåé äëèíû òåëà
Ma l e s
0 0.268 0.207 0.220 0.303 0.475 0.444 0.525 0.161
0.1 0.243 0.219 0.187 0.354 0.462 0.385 0.538 0.156
1 0.235 0.233 0.242 0.286 0.470 0.479 0.529 0.138
2 0.215 0.243 0.261 0.296 0.452 0.485 0.548 0.149
3 0.220 0.235 0.225 0.316 0.461 0.460 0.538 0.145
3–20 0.224 0.230 0.226 0.308 0.458 0.470 0.542 0.143
8–20 0.219 0.233 0.246 0.302 0.459 0.484 0.541 0.144
Fema l e s
0 0.264 0.192 0.241 0.305 0.456 0.431 0.544 0.164
0.1 0.267 – – 0.306 – 0.426 – –
1 0.231 0.240 0.219 0.308 0.471 0.462 0.529 0.135
2 0.229 0.260 0.187 0.310 0.488 0.457 0.512 0.134
3 0.221 0.253 0.250 0.312 0.458 0.475 0.526 0.136
3–20 0.212 0.241 0.219 0.327 0.447 0.463 0.549 0.130
8–20 0.210 0.245 0.232 0.320 0.447 0.469 0.553 0.127
Age (years) Sector I Sector II Sector III Sector IV Sector I+II Sector
II+III
Sector
III+IV
Lpect
Sector III demonstrates the most intricate growth pattern. The length of this sec-
tor least correlates with the total body length in both sexes. The coefficient estimates in
the allometry equations are rather rough. This is the only sector characterized by the
tendency of sexual differences in growth rate. In general, Sector III demonstrates po-
sitive allometry in males in the course of growth, taking 22% of total body length in
neonates and 24.2% in specimens at age of 1 year. In adult males this index remains
constant on an average, about 24%. In females the allometry is negative in juveniles,
and in adults this measurement does not correlate with the body length at all. The index
decreases from 24% in neonates to 23.2% in specimens at the age of 8–10 years. At the
same time, no sexual differences were found in the absolute length of Sector III.
The length of the basis of the dorsal fin, which is located within Sector III, demon-
strates slightly negative allometry and does not differ by growth rate in the sexes. The
sexual differences in allometry of dorsal fin height are not significant.
Therefore, it follows that the sexual differences in growth rates concern mostly the
posterior part of Sector III. In particular, this explains the outwardly paradoxical fact
that b coefficient in the allometry equation for the distance from the tip of the rostrum
to the centre of the genital slit (measurement 7) in females is somewhat greater than
for the distance to the anus (measurement 8). However, the data by A. J. Read and
K. A. Tolley (1997) do not confirm such a tendency.
The rapid early growth of Sector III in males is explained by the necessity for the
development of large genitalia, which are the specific character for harbour porpoise. The
testes weight in this species can reach to 4% of the body weight (Read, 1990). Due to
the comparatively small body size of the animals from the Sea of Azov and the Black Sea,
this tendency can be exhibited in them more clearly than in porpoises from other regions.
Sector IV demonstrates isometry or slightly positive allometry in growth of both
sexes. The index of this sector changes on an average from 30.2 to 30.5% in males, from
30.8 to 32% in females. In the majority of adult animals of a given sex the lengths of
Sector IV are approximately equal; they do not depend on the total body length.
However, in the longest females the absolute size and index of this sector increase, the
latter up to 35–41%. Sector IV is characterized by stable growth of the absolute size at
the age of 1–4 years, when the growth of the other body parts already ceases. Thus,
Sector IV, as well as Sector II, determines the total body length.
A. Galatius and C. C. Kinze (2003) report the partial ankylosis in the forepart of
the caudal department of the vertebral column in males begins at the age of more than
3 years and in females at the age of more than 4 years. Approximately at the same time
the growth of Sector IV ceases, and the further increment of the total body length
becomes very insignificant.
The length of Sector IV shows a strong correlation with the total body length. In
its growth, like in the growth of the whole organism, two stages can be distinguished,
which are best demonstrated in the growth of males from the Sea of Azov. The coeffi-
cients of determination r2 of the two-stages growth equations (the Gompertz formula)
for Sector IV are significantly higher in males than those of the one-stage equation. At
the same time, there are no significant differences in the coefficients of determination
of the growth equations for the rest of the body (sum of Sectors I-III). Thus, it is in
the growth of Sector IV where the periodicity is exhibited. The shift of the growth stages
takes place at the age between 1.5 and 2 years in males; in females this regularity is not
so clearly expressed, as well as in the growth of the total body length (Gol’din, 2004).
Thus, it is reasonable to suggest that it is Sector IV that is responsible for exhibiting the
stages in the growth of the organism.
Sector IV contains a great mass of the skeletal musculature. It is the caudal part
that is the main driver of a cetacean’s body. Thus, the periodicity of the growth of
organism in the porpoise is determined by the growth regularities of the locomotor mus-
culoskeletal complex.
63Body proportions of harbour porpoise Phocoena phocoena
A remarkable characteristics in the pattern of growth of Sector IV is that the peri-
odicity of the growth does not reveal itself in any allometry changes. This phenomenon
is explained by a very strong correlation of the length of Sector IV and total body length
during all the life span.
The maximum length of Sector IV was observed in the largest animals. That is why
studying the populations of harbour porpoises of the largest body size — in the waters
off Pyrenean peninsula, Africa, Pacific Ocean — is a subject of special interest.
The distance from the tip of the rostrum to the anus (measurement 8), including
the length of Sectors I–III, demonstrates slightly negative allometry or isometry (the
differences between the b coefficient and 1 are insignificant). The slightly negative
allometry is also characteristic for measurement 6 (the distance from the tip of the ros-
trum to the posterior edge of the dorsal fin basis) (the differences between the b coef-
ficient and 1 are significant).
The allometry of the posterior part of the porpoise body was studied in detail by
Van Utrecht (1978). According to his study, all the measurements in Sectors III and
IV in both sexes are characterized by isometry.
Almost all the measurements of the flippers and tail flukes are characterized by
negative allometry. Similar regularities were reported by other authors (Zalkin, 1938;
Van Utrecht, 1978; Read, Tolley, 1997; Galatius, personal communication).
The measurements of the dorsal fin and tail flukes (except the flukes span), as well
as the head measurements, are characterized by low correlation with the total body length.
Negative allometry and low correlation with total body size are observed in struc-
tures early developing and undergoing high pressures since the moment of birth: head
and fins. The lengths of Sector I and flippers better correlate with the total body length,
so the growth of these structures should last longer.
When comparing the lengths of pairwise summed sectors (I+II, II+III, III+IV) it
appears that only the middle body part (Sectors II+III) shows stable positive allometry
in both sexes — both during the growth period and in adult animals. The index of this
part increases in 4% during the lifespan in both sexes. However, two sectors, II and III,
contribute to the increment in males, while in females it is only Sector II, which at the
same time compensates the decrease of the portion of Sector III. Changes in the ratio
of the anterior and posterior parts of the body, mentioned by all the authors who stu-
died allometry in harbour porpoise (Zalkin, 1938; Van Utrecht, 1978; Stuart, Morejohn,
1980; Read, Tolley, 1997), are also observed in our sample, yet they appear less signif-
icant. The allometry equations for the pairwise summed sectors are exactly equal in both
sexes (the coefficients are printed in bold type in table 1). Thus, every value of the body
length is characterized by a specific dynamic ratio of proportions determined by a strict
regularity. The functional meaning of this ratio requires further study.
Sexual dimorphism of absolute sizes is exhibited in the lengths of Sectors II and
IV, which are characterized by isometry or positive allometry in both sexes. These sec-
tors cause sexual dimorphism in the total body length. The mean length of these sec-
tors starts to differ in males and females since the age of 2 years. Among sexually mature
specimens the length of Sector II in females is 4 cm more and the length of Sector IV
is 6 cm more than in males. However, due to high variability the differences in the
length of Sector II do not reach the 95% confidence level.
The position of the genital slit is another apparent and repeatedly reported inter-
sexual difference in harbour porpoises. In neonates the slit, with the anus at its caudal
edge, is located in the posterior part of Sector III. Soon after birth, along with the
growth of Sector III, the genital slit in males gets displaced forward, to the anterior
edge of this sector. By the end of the first year of life the distance from the centre of
the genital slit to the anus increases from 2–3 cm to 12–22 cm and can reach 32 cm
in adult males. In females, the length of the genital slit increases with age but its loca-
64 P. E. Gol'din
tion does not change. This fact accounts for significant sexual differences in the allom-
etry equations for measurements 7 and 9.
Conclusions
1. On the basis of studies of the allometric growth of the body parts it was first
shown that the growth of body parts during the postnatal ontogenesis of harbour por-
poise differs substantially in four body sectors corresponding to the skull and depart-
ments of vertebral column. The anterior part and fins — the structures undergoing high
pressures since the moment of birth — are characterized by negative allometry, and
many of their measurements have low correlation with the total body size. The highest
rates of allometric growth are observed in the thoracic and caudal departments.
2. The obtained results have allowed to hypothesize that the two-stage growth of
the organism is determined by the periodicity of growth of the caudal department and
the locomotor musculoskeletal complex associated with it.
3. The anterior, posterior and partly overlapping them middle body parts are inter-
connected by a ratio of proportions expressed by allometry equations and not depen-
ding on sex. The middle part of the body is characterized by maximum rates of allom-
etry in both sexes.
4. Sexual dimorphism of the body size is determined by body parts, which exhib-
it positive allometry or isometry in both sexes. Sexual differences in growth rates are
observed in the lumbar department, which grows faster in males. However, these dif-
ferences do not influence the absolute size except the anus-genital distance.
The author sincerely thanks the staff of the Department of Zoology of V. I. Vernadsky Taurida National
University (Simferopol), for the assistance during this study; D. V. Markov for many years of help in field
expeditions; V. V. Pavlov for the kindly provided data; À. Galatius, G. A. Klevezal’, C. H. Lockyer, K. Tolley
for their participation in the discussion of the results; M. V. Yurakhno for his valuable remarks concerning
the manuscript. This study was partially supported by the Award of President of Ukraine for the Graduate
Students and the Grant-in-Aid of the Society for Marine Mammalogy.
Galatius A., Kinze C. C. Ankylosis patterns in the post-cranial skeleton and hyoid bones of the harbour
porpoise (Phocoena phocoena) in the Baltic and North Sea // Can. J. Zool. — 2003. — 81, N 11. —
P. 1851–1861.
Gol’din P. E. Bone of lower jaw of harbour porpoise (Phocoena phocoena relicta Abel, 1905) as a registering
structure [Ãîëüäèí Ï. Å. Êîñòü íèæíåé ÷åëþñòè àçîâêè (Phocoena phocoena relicta Abel, 1905)
êàê ðåãèñòðèðóþùàÿ ñòðóêòóðà] // Uchenye Zapiski Tavricheskogo Nationalnogo Universiteta. Ser.
Biologiya. — 2003. — 16 (55), N 2. — P. 61–69. — Russian.
Gol’din P. E. Growth and body size of harbour porpoise (Phocoena phocoena) (Cetacea, Phocoenidae) in
the Sea of Azov and the Black Sea // Vestnik Zoologii. — 2004. — 38, N 4. — P. 59–73.
Read A. J. Reproductive seasonality in harbour porpoises from the Bay of Fundy // Can. J. Zool. — 1990. —
68, N 2. — P. 284–288.
Read A. J., Tolley K. A. Postnatal growth and allometry of harbor porpoises from the Bay of Fundy //
Can. J. Zool. — 1997. — 75, N 1. — P. 122–130.
Stuart L. J., Morejohn G. V. Developmental patterns in osteology and external morphology in Phocoena
phocoena // Age determination of toothed whales and sirenians. Rep. Int. Whal. Commn. (Special
issue 3). — Cambridge : IWC, 1980. — P. 133–153.
Van Utrecht W. L. Age and growth in Phocoena phocoena Linnaeus, 1758 (Cetacea, Odontoceti) from the
North Sea // Bijdr. Dierkd. — 1978. — 48, N 1. — P. 16–28.
Zalkin V. I. Morphological characteristics, taxonomic status and zoogeographical position of harbour porpoise
from the Sea of Azov and the Black Sea [Öàëêèí Â. È. Ìîðôîëîãè÷åñêàÿ õàðàêòåðèñòèêà,
ñèñòåìàòè÷åñêîå ïîëîæåíèå è çîîãåîãðàôè÷åñêîå çíà÷åíèå ìîðñêîé ñâèíüè Àçîâñêîãî è
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65Body proportions of harbour porpoise Phocoena phocoena
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| id | nasplib_isofts_kiev_ua-123456789-3385 |
| institution | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| issn | 0084-5604 |
| language | English |
| last_indexed | 2025-12-07T13:13:15Z |
| publishDate | 2005 |
| publisher | Інститут зоології ім. І. І. Шмальгаузена НАН України |
| record_format | dspace |
| spelling | Gol'din, P. E. 2009-07-07T10:04:16Z 2009-07-07T10:04:16Z 2005 Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea / P. E. Gol’din // Вестн. зоологии. — 2005. — Т. 39, № 5. — С. 59–65. — англ. 0084-5604 https://nasplib.isofts.kiev.ua/handle/123456789/3385 591.4:599.536(262.5) Исследования относительного роста частей тела морской свиньи впервые показали, что он в постнатальном онтогенезе существенно различается в четырех секторах, соответствующих черепу и отделам позвоночника. Наибольшими темпами относительного роста отличаются грудной и хвостовой отделы. Средняя часть тела отличается наибольшими темпами относительного роста у обоих полов. Выдвинута гипотеза о том, что двухстадийный рост организма определяется периодичностью роста хвостового отдела. Половой диморфизм размеров тела создается за счет частей тела, проявляющих положительную аллометрию или изометрию у обоих полов, однако различия в абсолютных величинах затрагивают лишь анально-генитальное расстояние. The studies of the allometric growth of the body parts in harbour porpoises have shown that the growth of body parts during the postnatal ontogenesis of the harbour porpoise differs substantially in four body sectors, which correspond to the skull and the parts of vertebral column. The highest rates of allometric growth are observed in the thoracic and caudal departments. The middle part of the body is characterized by maximum rates of allometry in both sexes. The hypothisis was brought forward that the two-stage growth of the organism is determined by the periodicity of growth of the caudal department. Sexual dimorphism of the body size is determined by the body parts, which exhibit positive allometry or isometry in both part. Sexual differences in growth rates are observed in the lumbar part but the differences of absolute length values concern only anusgenital distance. en Інститут зоології ім. І. І. Шмальгаузена НАН України Морфология Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea Пропорции тела морской свиньи Phocoena phocoena (Cetacea, Phocoenidae) в Азовском и Черном морях Article published earlier |
| spellingShingle | Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea Gol'din, P. E. Морфология |
| title | Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea |
| title_alt | Пропорции тела морской свиньи Phocoena phocoena (Cetacea, Phocoenidae) в Азовском и Черном морях |
| title_full | Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea |
| title_fullStr | Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea |
| title_full_unstemmed | Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea |
| title_short | Body Proportions of Harbour Porpoise Phocoena phocoena (Cetacea, Phocoenidae) in the Sea of Azov and the Black Sea |
| title_sort | body proportions of harbour porpoise phocoena phocoena (cetacea, phocoenidae) in the sea of azov and the black sea |
| topic | Морфология |
| topic_facet | Морфология |
| url | https://nasplib.isofts.kiev.ua/handle/123456789/3385 |
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