Comparing the growth and weight relationships of the Arctic toothed whales (Monodontidae)

This update on the small cetacean body mass database describes what I’ve accumulated for the narwhal (Monodon monoceros) and the beluga (Delphinapterus leucas). Like my previous posts on river dolphins and kogiids, I will also provide some insights into my methods of data collection and discussion the nature of some of the information I’ve collected.

The Horned and the White Whales

Both the beluga and narwhal are medium-sized toothed whales that live along the Arctic and subarctic waters. Belugas possess a circumpolar distribution whereas the narwhal is only native to the Northern Europe and Eastern Canada, with few vagrants observed off Alaska. The first scientific description of the beluga came from observations of harvests along the Ob river and was assigned the name Delphinus leucas (Pallas, 1776), meaning “white dolphin’’. The current generic name, Delphinapterus (La Cépède, 1804), refers to the absence of the dorsal fin. Having been formally described in Russia, the common name is derived from the word for ‘’white’’.

Before being formally described, the narwhal’s tusk was well known from its association with the Biblical unicorn (Bartholin, 1678; Pluskowski, 2004; Tomilin, 1967; Worm, 1655). The narwhal was also among the first whales described in System Naturae (Linnaeus, 1758). The tusk is referenced in both the generic (“One tooth’’) and specific names (“one horn’’).  The common name is thought to have come from the Old Norse term meaning ‘’corpse whale’’ (Mansfield et al., 1975), in reference to the mottled skin resembling a decayed appearance.

While early taxonomic schemes classified narwhals and belugas within the Delphinidae family along with oceanic dolphins (Kleinenberg et al., 1969), they have since been separated into their own family, Monodontidae. Fossil species firmly classified into this family such as Bohaskaia monodontoides in the Northwest Atlantic (Vélez-Juarbe & Pyenson, 2012), Haborodelphis japonicus in the Northwest Pacific (Ichishima et al., 2019) and Casatia thermophila  from the Mediterranean (Bianucci et al., 2019; Merella et al., 2022) were from the early Pliocene. The latter supports that Monodontids evolved in more temperature waters prior to cooling evens (Bianucci et al., 2019). The oldest, Denebola brachycephala, from the late Miocene (~11-5.4 million years ago) was found in the Eastern North Pacific (Barnes, 1984).  

The unfused cervical vertebrae distinguishes extant Monodontidae from all other cetaceans. This feature, along with others, supports that Odobenocetops is related to monodontids (de Muizon, 1993; de Muizon et al., 1999; de Muizon & Domning, 2002). Molecular studies indicate that monodontids are the nearest extant relatives to porpoises (Phocoenidae). While narwhals and belugas are classified into separate genera, hybridization has been confirmed (Heide‐Jørgensen & Reeves, 1993; Skovrind et al., 2019).

Indigenous populations along North America, Russia, and Greenland all have long histories of harvesting both species (Hay, 1984; Kleinenberg et al., 1969), which is currently regulated by each nation’s government (Gonzalez, 2001; Heide-Jørgensen et al., 2010; Huntington & The Communities of Buckland, 1999; Shpak et al., 2020; Tyrrell, 2008) rather than the IWC. It is from these harvests that we obtain much of our recent data on these species (Garde et al., 2007; Hansen et al., 1990; Suydam, 2009).

While narwhals have rarely ever been held in captivity, belugas were the species to first, with P.T. Barnum being one of the earliest proprietors (Ceta-Base: Online Marine Mammal Inventory, 2010).

Life History and Growth

Tables 1 & 2 compares the size parameters for different populations of both species. Belugas are known to change in pigment with age. They’re an off-black color at birth and progressively become whiter towards adulthood (Kleinenberg et al., 1969). This was a useful, but imprecise indicator of age for the earliest examinations into this species’ life history. The transition towards a full-white pigment appears to closely coincide with sexual maturity in males, but females typically mature before becoming fully white (Kleinenberg et al., 1969).

Age estimations in both species are conventionally calculated using laminations from growth layers in the teeth (Mansfield et al., 1975; Sergeant & Brodie, 1969). In the case of the narwhal, this is done with the embedded incisor opposite of the tusk. There’s some debates as whether 1 or 2 layers form per year for the beluga (Luque & Ferguson, 2010; Sergeant, 1973), however I’m leaning towards the 2 layer/year conversion based on validation from captive individuals of known age (Brodie & Haulena, 2018). By this standard, belugas around the world appear become sexually mature at consistent ages of around 5 and 7-9 years for males and females respectively (Brodie, 1971; Kleinenberg et al., 1969; Sergeant, 1973; Suydam, 2009). Physical maturity generally appears to occur between 10-20 years of age.

Use of another technique called aspartic acid racemization (AAR), an aging technique applied to the eye lens (Garde et al., 2007), suggests narwhals also exhibit biannual deposition rate for growth layers. The maximum ages I could find for both species appears to exceed 50 years for belugas (Sergeant, 1973) and 107.7 years for narwhals (Garde et al., 2022).

Table 1 : Size parameters for D. leucas

RegionLb (cm)Lsm (cm) M/FL(cm) M/FLmax  M/FSource
W. Hudson Bay151310 / 290377 / 328448 / 400(Doan & Douglas, 1953; Doidge, 1990b; Sergeant, 1973)
E. Hudson Bay316 / 280a349 / 330440b / 400b(Doidge, 1990b, 1990a)
Cumberland Sound160378c / 308427 / 362450b / 400b(Brodie, 1971; Luque & Ferguson, 2010)
White & Kara Seas160380d / 350438 / 378479 / 420(Heide-Jørgensen & Teilmann, 1994; Kleinenberg et al., 1969; Ognetov, 2007)
Alaska155369e / 289432f / 370f485 / 410(Burns & Seaman, 1986; Suydam, 2009)
St. Lawrence Estuary160381g / 336g416 / 365456 / 409(Lair et al., 2015; Lesage et al., 2014; Vladykov, 1944)
W. Greenland155a390 / 345483 / 386545 / 482(Degerbøl & Nielsen, 1930; Heide-Jørgensen & Teilmann, 1994)
Okhotsk Sea170390h / 343ij / 390590a / 490a(Dorofeev & Klumov, 1936; Kleinenberg et al., 1969)
a. Mean of listed range or binned value
b. Approximation from figure.
c. Estimation from Gompertz curve from Luque & Ferguson, 2010 at age 7
d. Approximation from Ognetov, 2007’s age-length key for age 8
e. Average of Eastern Chukchi and Cook Inlet Males at 7 years (Suydam, 2009).
f. Average of multiple regions across Alaska (Suydam, 2009).
g. Estimation from growth curve from Lesage et al., 2014.
h. Based on approximate mean length from transition from grey to white (Dorofeev & Klumov, 1936).
i. Mean of smallest mature female and largest maturing female (Kleinenberg et al., 1969).
j. Insufficient data, likely > 470 cm based on modal length of white males (Dorofeev & Klumov, 1936).

Belugas are evidently a polymorphic species with respect to adult size. The Hudson Bay populations are the smallest, West Greenland and Okhotsk Sea belugas are the largest, and all other regions being intermediate. The largest belugas to be reliably documented were three females with binned lengths of 480-499 cm and two males 580-599 cm from the Okhotsk Sea (Dorofeev & Klumov, 1936).

Table 2: Size parameters for M. monoceros

RegionLb (cm)Lsm (cm) M/FL(cm) M/FLmax  M/FSource
Baffin Island161395 / 340470 / 415470 / 415(Hay, 1984; Hay & Mansfield, 1989; Mansfield et al., 1975)
W. Greenland456 / 399  560 / 490(Garde et al., 2015)
E. Greenland    462 / 405480 / 441(Garde et al., 2015)
SE. Greenlanda  463 / 406501 / 441(Garde et al., 2022)
a. Data includes some individuals with East Greenland Sample from Garde et al., 2015

By contrast, the adult size of narwhals appears more consistent between Eastern Canada and Greenland. A 560 cm male and 490 cm female (Garde et al., 2015) gives credence to early literature assigning maximum lengths of 600 cm for males and 500 cm for females (Tomilin, 1967). The typical size of narwhals corresponds to that of the larger-sized beluga populations. The tusk of males narwhals typically grows to 160-200 cm (Garde et al., 2015, 2022). Eighteenth century catch logs documents lengths of 267 cm (Mitchell & Reeves, 1981) and one manuscript states that the narwhal’s tusk reach 315 cm (Bruemmer, 1971). I was not able to obtain the latter source to verify the context of this measurement for myself. Excluding both of these, there are at least two verified records of tusks reaching 240 cm (Beasley, 1904; Garde et al., 2015).

Body weight

Figures 1 and 2 below are the plots for the weight data I’ve accumulated for the beluga (n= 347) and narwhal (n = 96), respectively. Nearly all were from literature as only a measly single individual was included from the USNM database. I elaborate on the heaviest individuals while describing the Okhotsk Sea material (Govorkov, 1934), but for now I’ll say the heaviest masses recorded outside of this region for each sex was a 465 cm male from the Kara Sea that weighed 1,480 kg (Ognetov, 2007) and a 388 cm female from St. Lawrence that weighed 1,003 kg (Lair et al., 2015). The corresponding records for narwhals were a 470 cm / 1,789 kg male and a 410 cm / 1,148 kg female from West Greenland (Garde et al., 2007).

Figure 1: Weight data for D. leucas

In the regional breakdown for belugas, the St. Lawerence region was by far the most numerous (n = 205), with the next two most common regions being the Hudson Bay (n = 45) and those from the White, Kara, and Barents Seas (n = 40). I even had a small sample of captive individuals (n=10), which includes one large male that weighed 1,200 kg (Brodie et al., 2013). The dataset covers nearly all the major regions except for Alaska and West Greenland. For narwhals, it was an even split between Baffin Island (n = 46) and Greenland (n = 50).

Figure 2: Weight data for M. monoceros

The narwhal dataset excludes four underweight outliers, two of which were from strandings. The weights of 16 eviscerate belugas from the West Hudson Bay were excluded (Doan & Douglas, 1953), as a previous study (Doidge, 1990b) has shown that  incorrect assumptions on correction factors confound apparent differences in weight relationships (Sergeant & Brodie, 1969). An incomplete weighing for a 1,000+ kg female was also excluded (Doidge, 1990b).

Validating Govorkov’s data

My biggest goal when compiling the data was to find a decent sample from the Okhotsk Sea. Unfortunately, it appears nearly all collection of such material was restricted to obscure Soviet research from the early-mid 20th century (Kleinenberg et al., 1969). There are currently some sources I have not been able to access (Arsenʹev, 1935, 1939), but was mentioned in other literature (Heptner et al., 1988; Kleinenberg et al., 1969; Tomilin, 1967; Vladykov, 1944). I was fortunately able to find one source (Govorkov, 1934), however this has been flagged as potentially unreliable by other authors (Kleinenberg et al., 1969; Tomilin, 1967) as it reports males as large as 667 cm and females reaching 651 cm. Figure 3 shows the diagram of the measuring procedure Govorkov presents.

Figure 3: Reported measuring scheme from Govorkov, 1934

  Two things immediately struck my attention upon reviewing the source myself. First, was that the total length, as presented in the diagram, was intended to be the zoological standard. The other was that the 667 cm male and 651 cm female were the only two that exceeded 600 cm. I wouldn’t expect such a discontinuity of over 0.5 m for the next largest whales if the entire sample was equally biased. Since fluke widths, a less ambiguous measurement, was also recorded for each whale. This allowed for comparisons with other samples to validate the reliability of total lengths (Figure 4) as performed in other studies (Doidge, 1990b; Heide-Jørgensen & Teilmann, 1994). Due to the likelihood of morphological differences between geographically distant populations, I restrict my analysis to measurements taken from 31 belugas from the Amur Estuary and Yenisei Bay in the Far East (Arsenʹev, 1936; Drukker & Gakichko, 1936), as they were the nearest population. Due to size differences between the samples, I only compared Okhotsk belugas as large as 410 cm.

Figure 4: Fluke width comparisons between Okhotsk Sea with Far East Belugas

AIC model selection slightly preferred pooling the two samples (weight = 0.61) and it can be seen graphically in Figure 4 that there’s no obvious difference between the sources. It seems that the length measurements for belugas up to 410 cm showed no obvious signs of biased length measurements. Table 3 compares Govorkov’s sample against regression predictions for TL from a large sample of belugas sampled in the Eastern Chukchi sea (Suydam, 2009) and West Greenland (Heide-Jørgensen & Teilmann, 1994), TL’s for males appear accurate up to 576 cm, excluding only the 667 cm male. Meanwhile, TLs for females appears becomes uncertain past 410 cm. I suspect females with estimated TLs exceeding 500 cm may potentially have been misidentified males. Table 3 lists individuals in descending order of length and boundaries of reliability are highlighted.

Table 3: Fluke width and estimated TL from Govorkov, 1934

SexTotal length (cm)Fluke span (cm)Mass (kg)Estimated TL-SuydamEstimated TL-Heide-Jorgensen & Teillman, 1994
F6511571823643.88661.2173913
F5841251213586.4522.0869565
F5611321728621.4552.5217391
F5241121493521.4465.5652174
F490981096451.4404.6956522
F45490946411.4369.9130435
F41088873401.4361.2173913
F401a661312291.4265.5652174
F35076848341.4309.0434783
F32075720336.4304.6956522
F31566670291.4265.5652174
F30448720201.4187.3043478
F30063488276.4252.5217391
F23048248201.4187.3043478
F22646224191.4178.6086957
F19844215181.4169.9130435
M6671361412559.88569.9130435
M5761511956619.88635.1304348
M5361291632531.88539.4782609
M463971437403.88400.3478261
M4481041347431.88430.7826087
M440701593295.88282.9565217
M401102816423.88422.0869565
M39979926331.88322.0869565
M37574742311.88300.3478261
M36278875327.88317.7391304
M22545448195.88174.2608696
M19045202195.88174.2608696
M17039194171.88148.173913
M15338160167.88143.826087
a. While much larger than estimated TL, the corresponding weight suggests that the reported TL is accurate, and this female is simply at the extreme end of proportions.

While fluke measurements provided promising inferences, I didn’t stop my assessment there, as I also employed a mass-prediction model to validate the length measurements against their weight data. The text explains that girth measurements along the dorsal side were taken (probably only 60-80% of the full girth). Total length and maximum girth can potentially describe nearly all the variation in weight in small cetaceans (Garde et al., 2022; Lockyer, 1993). Therefore, overestimation in total length or girth can be detected if the estimated mass greatly exceeds the reported mass. Since Govorkov’s sample records partial girths, we should expect the estimated masses to lie consistently below the reported mass.

Table 4: Real and estimated masses from Govorkov, 1934

SexLength (cm)Partial girth (cm)Mass (kg)Estimated mass (kg)
F65118418232238.212919
F58423712132136.088562
F56126017282097.900223
F52423814931694.167622
F49021010961334.466323
F4541809461008.317498
F410170873774.7451589
F4011351312621.8051084
F350204848629.8911237
F320168720448.8339329
F315201670495.8081532
F304178720419.3027936
F300165488385.0483043
F230162248213.5592729
F226156224199.8901635
F198115215a119.5678005
M66736714123946.653259
M5762701956b2284.723283
M53615616321298.7888
M46319614371121.076193
M44822613471160.751219
M44019615931003.882937
M401190816802.1795509
M399178926755.873231
M375176742655.2530046
M362234875750.5864558
M225155448a,c197.0318359
M190116202a110.0530961
M170113194a84.80956762
M15394160a58.84386526
a. All weights are twice as much as expected. This pattern is only seen in young animals, suggesting these were measured using different equipment that measured weight in lbs. as in Vladykov, 1944.
b. Given the consistency with fluke width, the girth was likely overestimated rather than TL.
c. Female of near-exact same dimensions weighed half as much. Converting to kg removes any difference.

A previously published mass prediction model that uses the girth near the umbilicus (Doidge, 1990b) is not appropriate for such a comparison, as the girths reported by Govorkov were at the axilla, as taken in larger whales (George, 2009; Rice & Wolman, 1971; Víkingsson et al., 1988). Therefore, I fitted my own model using a sample of 12 belugas from other data that measured the axillary girth (Drukker & Gakichko, 1936; Vladykov, 1944). This model had a notably higher correlation (r2= 0.991) than other models for belugas and narwhals that used umbilical girths (Doidge, 1990b; Garde et al., 2022).

0.00003672 (Length in cm)2.167 (Girth in cm)0.7453

Remarkably, Table 4 shows that overestimation in mass coincides almost exactly where we see a loss in consistency in TL estimation from fluke width regressions in both sexes. Furthermore, there’s compelling evidence of a unit conversion error for most of the small individuals (< 230 cm). Since this is only observed in calves, they were likely measured using a separate set of equipment that measured in lbs. that weren’t converted to kilograms. Dividing these values by 2.2 removes this discrepancy.

It appears clear to me now that Govorkov’s data was likely pooled by different direct sources or was collected under variable conditions that impacted reliability. Cross-referencing Tables 3 & 4 supports that 23 of the TL measurements were reliable, with a 576 cm male weighing 1,956 kg and a 401 cm female weighing 1,312 kg being the heaviest records for the species.

The 651 cm female was excluded altogether, despite estimated TLs being consistent with its reported length, as it’s such an extreme outlier, even if sex was misidentified. A female reported as 454 cm and 946 kg was a bit of a judgement call, but opted to slightly downsize the length using the fluke width regression for females from the Eastern Chukchi Sea (Suydam, 2009). As for the remaining five, I chose to downsize their lengths using the unisex formula from the West Greenland population (Heide-Jørgensen & Teilmann, 1994) as it seemed better suited for both large specimens and those of uncertain sex.

Figure 1 shows that all individuals exceeding 500 cm appear proportionately lean, this seems consistent with a 520 cm beluga reported to weigh 1,372 kg (Mikhalev, 2019). It’s likely that exceptionally long individuals are leaner than those that are only slightly above average.

Weight relationships.

When looking closely at the logarithmic plots in Figures 1 & 2, there are some discontinuities to the linear pattern between juveniles and the largest individuals, causing the overall trend to appear wavy. This is particularly clear in the narwhal data. The discontinuity for individuals below 290 cm was observed for the weight data from the St. Lawrence area population (Larrat & Lair, 2022). This roughly corresponds with existing data on the size of weaning at about 230-280 cm (Kleinenberg et al., 1969; Sergeant, 1973; Suydam, 2009) and appears to apply to narwhals as well. The discontinuity for large individuals appears to be due to the accumulation of blubber in older males after their growth ceases (Hay, 1984). This pattern was also observed in other species in my database, including the 700 kg kogiids noted in my last post.

Not accounting for at least the discontinuity between juveniles and maturing individuals will confound AIC model comparisons. After separating the datasets for both species into two size classes (above 290 cm and below 290 cm), AIC model selection revealed that the narwhal’s weight relationship exhibited notable sexual differences (delta AIC > 2) and weak regional differences while the opposite was found for belugas. Both patterns were more pronounced for individuals > 290 cm. Table 5 provides a list of linear regression parameters obtained from the current dataset and those obtained from literature. All belugas exceeding 500 cm were excluded from regressions for adult-sized individuals.

Table 5: Power function parameters for Monodontidae

SpeciesSexRegiona*br2NSource
M. monocerosallBaffin Island29.52  2.48038 41Hay, 1984
M. monocerosMalePooled29.262.5570.96524Current post
M. monocerosFemalePooled32.32.4030.97871Current post
D. leucasa  BothHudson Bay16.872.536 16Sergeant & Brodie, 1969
D. leucasBothSt. Lawrence25.292.605 10Sergeant & Brodie, 1969
D. leucasBothHudson Bay20.892.580.9236Doidge, 1990 a & b
D. leucasMaleW. Greenland25.692.47 41Heide-Jorgensen & Tillman, 1994
D. leucasFemaleW. Greenland35.982.21 57Heide-Jorgensen & Tillman, 1994
D. leucasBothW. Greenland28.252.37 98Heide-Jorgensen & Tillman, 1994
D. leucasMaleNorth Quebec10.993.37 20Heide-Jorgensen & Tillman, 1994
D. leucasFemaleNorth Quebec40.362.03 16Heide-Jorgensen & Tillman, 1994
D. leucasBothNorth Quebec15.542.85 36Heide-Jorgensen & Tillman, 1994
D. leucasBothCaptive40.332.080.949bRobeck et al., 2005
D. leucasMaleCaptive40.842.132 5bRobeck et al., 2005
D. leucasFemaleCaptive47.502 4bRobeck et al., 2005
D. leucasBothPooled21.202.6470.935179Mikhalev, 2019
D. leucasMaleWhite & Kara Seas21.382.6789 140Ognetov, 2007
D. leucasFemaleWhite & Kara Seas23.152.5668 156Ognetov, 2007
D. leucasBothWhite & Kara Seas21.912.647 296Ognetov, 2007
D. leucasBothPooled18.742.7730.959342Current Post
D. leucasBothCaptive18.702.7810.97010Current Post
D. leucasBothHudson Bay26.432.4130.89645Current Post
D. leucasBothArctic Route20.472.6760.97850Current Post
D. leucasBothOkhotsk Sea19.562.8860.96224Current Post
D. leucasBothSt. Lawrence17.112.8450.962205Current Post
*Constant of proportionality converted so length is in meters and mass is in kg.
a. Uses data from eviscerated whales, is very negatively biased.
b. Represents number of separate individuals; sample is derived from unspecified number of longitudinal measurements.

While AIC modeling suggests there’s a significant difference between the weight relationships between the two species, overall, there’s still decent similarity, as to be expected for close relatives (Figure 5). A general, pooled weight relationship of the total dataset is expressed below.

Mass (kg) = 20.41 (Length, meters)2.728

Figure 5: Comparison between mass data for M. monoceros and D. leucas.

Maximum possible weight

Despite having covered a lot, there’s still one big dataset I have left. Using the weight model I developed earlier, I estimated the weight for 154 (excluding 3 emaciated outliers) St. Lawrence belugas captured during 1938-1939 (Vladykov, 1944). When compared to data collected from 1983-2012 (Lair et al., 2015), there seems to be less spread in the data for the estimated weights. There’s also no detectable discontinuity between juveniles and adults as found in the modern sample. These differences could be due to difference in sampling (commercial catch vs strandings), temporal differences in the condition of the stock, or due to the model not being sufficiently sensitive to variation in girth. The power function for the estimated masses of the 1938-1939 catch is as follows.

Mass (kg) = 19.40 (Length, meters)2.820

Figure 6: Comparison of weight data for St. Lawrence belugas

Girth in the heaviest males typically exceeds slightly over 70% of the body length. Using this against the maximum confirmed length of about 590 cm for male belugas from the Okhotsk Sea suggest that they could theoretically achieve a maximum weight of  3,300 kg. However, Govorkov’s data implies such a mass is never achieved, and the maximum mass may be closer to 2000-2500 kg.

What’s next?

After finishing off with my work with monodontids, I will likely be moving on to the blackfish species. That will likely take only another few weeks as I’m already certain I’ve cleared most of the data available within the literature. The current tally for the entire database is 3,208 from literature and 3,850 from other databases. Let’s see what the total looks like by the time the next post is up.

I continue to invite anyone to who would like to share any information they feel could be relevant to the email below

cetologyh@gmail.com

References

Arsenʹev, V. A. (1935). Some data on the Beluga (Delphinapterus leucas Pall.) and its Catch in Sakhalin. Far Eastern Fisheries, 13.

Arsenʹev, V. A. (1936). Морфологическая характеристика дальневосточной белухи [Morphological characteristics of the Far East Beluga]. Bulletin of the Far Eastern Branch of the Academy of Sciences of the USSR, 19, 131–144.

Arsenʹev, V. A. (1939). Distribution and Migration of the Far Eastern Beluga. Izvestiya TNIRO, 15.

Barnes, L. G. (1984). Fossil odontocetes (Mammalia: Cetacea) from the Almejas Formation, Isla Cedros, Mexico. PaleoBios, 42, 1–46.

Bartholin, T. (1678). Thomae Bartholini De unicornu observationes novae: Secunda editione auctiores & emendatiores editae a filio Casparo Bartholino. Apud H. Wetstenium.

Beasley, W. L. (1904). The Largest Narwhal Tusk. Scientific American, 91(15), 251–251. https://doi.org/10.1038/scientificamerican10081904-251a

Bianucci, G., Pesci, F., Collareta, A., & Tinelli, C. (2019). A new Monodontidae (Cetacea, Delphinoidea) from the lower Pliocene of Italy supports a warm-water origin for narwhals and white whales. Journal of Vertebrate Paleontology, 39(3), e1645148. https://doi.org/10.1080/02724634.2019.1645148

Brodie, P. F. (1971). A Reconsideration of Aspects of Growth, Reproduction, and Behavior of the White Whale (Delphinapterus leucas), with Reference to the Cumberland Sound, Baffin Island, Population. Journal of the Fisheries Research Board of Canada, 28(9), 1309–1318. https://doi.org/10.1139/f71-198

Brodie, P. F., & Haulena, M. (2018). Dentinal growth layer counts of captive, known-age, mother and daughter belugas (Delphinapterus leucas): Confirming two growth layer groups (GLG/2) per year; consequences for recovery and management. Journal of Cetacean Research and Management, 18, 23–31. https://doi.org/10.47536/jcrm.v18i1.420

Brodie, P. F., Ramirez, K., & Haulena, M. (2013). Growth and maturity of belugas (Delphinapterus leucas) in Cumberland Sound, Canada, and in captivity: Evidence for two growth layer groups (GLGs) per year in teeth. J. Cetacean Res. Manage., 13(1), Article 1. https://doi.org/10.47536/jcrm.v13i1.550

Bruemmer, F. (1971). Notes on sea mammals, Thule district, Greenland. Unpublished Manuscript, 29.

Burns, J. J., & Seaman, G. A. (1986). Investigations of belukha whales in coastal waters of western and northern alaska II. biology and ecology (pp. 223–357) [Technical Report]. Alaska Department of Fish and Game.

Ceta-Base: Online Marine Mammal Inventory. (2010). Captive Belugas: A Historical Record & Inventory (Europe, Canada, North America & United Kingdom). https://www.kimmela.org/wp-content/uploads/2012/09/captivebelugas_august2010.pdf

de Muizon, C. (1993). Walrus-like feeding adaptation in a new cetacean from the Pliocene of Peru. Nature, 365, 745–748. https://doi.org/10.1038/365745a0

de Muizon, C., & Domning, D. (2002). The anatomy of Odobenocetops (Delphinoidea, Mammalia), the walrus-like dolphin from the Pliocene of Peru and its palaeobiological implications. Zoological Journal of the Linnean Society, 134, 423–452. https://doi.org/10.1046/j.1096-3642.2002.00015.x

de Muizon, C., Domning, D. P., & Parrish, M. (1999). Dimorphic tusks and adaptive strategies in a new species of walrus-like dolphin (Odobenocetopsidae) from the Pliocene of Peru. Comptes Rendus de l’Académie Des Sciences – Series IIA – Earth and Planetary Science, 329(6), 449–455. https://doi.org/10.1016/S1251-8050(00)80070-1

Degerbøl, M., & Nielsen, N. L. (1930). Biologiske iagttagelser over og maalinger af hvidhvalen (Delphinapterus leucas (Pall.)) og dens fostre. Meddelelser Om Grønland, 77(3), 119–144.

Doan, K. H., & Douglas, C. W. (1953). Beluga of the Churchill Region of Hudson Bay (98; Fisheries Research Board of Canada Bulletin, p. 27).

Doidge, D. W. (1990a). Age and stage based analysis of the population dynamics of beluga whales, Delphinapterus leucas, with particular reference to the Northern Quebec population [Ph.D Thesis]. McGill University.

Doidge, D. W. (1990b). Age-length and length-weight comparisons in the beluga, Delphinapterus leucas. In T. G. Smith, D. J. S. Aubin, & J. R. Geraci (Eds.), Advances in Research on the Beluga Whale, Delphinapterus Leucas (pp. 59–68). Department of Fisheries and Oceans.

Dorofeev, S. W., & Klumov, S. K. (1936). К ВОПРОСУ ОБ ОПРЕДЕЛЕНИИ ВОЗРАСТА БЕЛУХИ И СОСТАВА КОСЯКОВ [About the age determination of the Beluga whale (Delphinaterus leucas Pall) and composition of the herd. Trudy VNIRO, 3, 24–34.

Drukker, G. F., & Gakichko, S. I. (1936). Pacific beluga whale as an industrial raw material. Trudy VNIRO, 3, 53–130.

Garde, E., Hansen, S. H., Ditlevsen, S., Tvermosegaard, K. B., Hansen, J., Harding, K. C., & Heide-Jørgensen, M. P. (2015). Life history parameters of narwhals (Monodon monoceros) from Greenland. Journal of Mammalogy, 96(4), Article 4. https://doi.org/10.1093/jmammal/gyv110

Garde, E., Heide-Jørgensen, M. P., Hansen, S. H., Nachman, G., & Forchhammer, M. C. (2007). Age-Specific Growth and Remarkable Longevity in Narwhals (Monodon monoceros) from West Greenland as Estimated by Aspartic Acid Racemization. Journal of Mammalogy, 88(1), Article 1. https://doi.org/10.1644/06-MAMM-A-056R.1

Garde, E., Tervo, O. M., Sinding, M.-H. S., Nielsen, N. H., Cornett, C., & Heide-Jørgensen, M. P. (2022). Biological parameters in a declining population of narwhals (Monodon monoceros) in Scoresby Sound, Southeast Greenland. Arctic Science, 8(2), 329–348. https://doi.org/10.1139/as-2021-0009

George, J. C. (2009). Growth, Morphology And Energetics Of Bowhead Whales (Balaena Mysticetus) [Thesis]. https://scholarworks.alaska.edu/handle/11122/9031

Gonzalez, N. (2001). Inuit Traditional Ecological Knowledge ofthe Hudson Bay Narwhal (Tuugaalik) Population. Department of Fisheries and Oceans.

Govorkov, I. V. (1934). Beluga: Trapping and Processing. Snabtekizdat.

Hansen, C., Nielsen, C., Dietz, R., & Hansen, M. (1990). Zinc, cadmium, mercury and selenium in minke whales, belugas and narwhals from West Greenland. Polar Biology, 10(7). https://doi.org/10.1007/BF00233702

Hay, K. A. (1984). The life history of the narwhal, Monodon monoceros l., in the eastern Canadian arctic [Ph.D Thesis, McGill University]. https://escholarship.mcgill.ca/concern/theses/v118rf31m

Hay, K. A., & Mansfield, A. W. (1989). Narwhal—Monodon monoceros (Linnaeus, 1758). In River dolphins and the larger toothed whales (Vol. 4, pp. 119–144). London ; New York : Academic Press. http://archive.org/details/riverdolphinslar00ridg

Heide-Jørgensen, M. P., Laidre, K. L., Burt, M. L., Borchers, D. L., Marques, T. A., Hansen, R. G., Rasmussen, M., & Fossette, S. (2010). Abundance of narwhals (Monodon monoceros) on the hunting grounds in Greenland. Journal of Mammalogy, 91(5), 1135–1151. https://doi.org/10.1644/09-MAMM-A-198.1

Heide‐Jørgensen, M. P., & Reeves, R. R. (1993). Description of an Anomalous Monodontid Skull from West Greenland: A Possible Hybrid? Marine Mammal Science, 9(3), 258–268. https://doi.org/10.1111/j.1748-7692.1993.tb00454.x

Heide-Jørgensen, M. P., & Teilmann, J. (1994). Growth, reproduction, age structure and feeding habits of white whales (Delphinapterus leucas) in West Greenland waters. Meddelelser Om Grønland, Bioscience, 39, 195–212.

Heptner, V. G., Nasimovich, A. A., Bannikov, A. G., & Hoffmann, R. S. (1988). Mammals of the Soviet Union (Vol. 2 part 3, pp. 1–1044). Smithsonian Institution Libraries and National Science Foundation. https://doi.org/10.5962/bhl.title.46297

Huntington, H. P., & The Communities of Buckland, E., Koyuk, Point Lay, and Shaktoolik. (1999). Traditional Knowledge of the Ecology of Beluga Whales (Delphinapterus lencas) in the Eastern Chukchi and Northern Bering Seas, Alaska. Arctic, 52(1), 49–61.

Ichishima, H., Furusawa, H., Tachibana, M., & Kimura, M. (2019). First monodontid cetacean (odontoceti, delphinoidea) from the early pliocene of the north‐western pacific ocean. Papers in Palaeontology, 5(2), 323–342. https://doi.org/10.1002/spp2.1244

Kleinenberg, S. E., Yablokov, A. V., Bel’kovich, B. M., & Tarasevich, M. N. (1969). Beluga (Delphinapterus leucas ): Investigation of the species. Israel Program of Scientific Translations.

La Cépède, B. G. de. (1804). Histoire naturelle des cétacées. Chez Plassan, Imprimeur-Libraire. https://doi.org/10.5962/bhl.title.4824

Lair, S., Gentes, M. L., Measures, L. N., & Michaud, R. (2015). Documentation de l’évolution du protocole d’examen des carcasses de béluga de l’estuaire du Saint-Laurent de 1983 à 2012 [Documentation of the evolution of the protocol for examination of beluga carcasses in the St. Lawrence Estuary from 1983 to 2012] (p. 65) [Technical Report]. Pêches et Océans Canada.

Larrat, S., & Lair, S. (2022). Body condition index in beluga whale (Delphinapterus leucas) carcasses derived from morphometric measurements. Marine Mammal Science, 38(1), 274–287. https://doi.org/10.1111/mms.12855

Lesage, V., Measures, L. N., Mosnier, A., Lair, S., Michaud, R., & Béland, P. (2014). Mortality patterns in St. Lawrence Estuary beluga (Delphinapterus leucas), inferred from the carcass recovery data, 1983-2012. Fisheries and Oceans Canada Canadian Science Advisory Secretariat.

Linnaeus, C. von. (1758). Caroli Linnaei…Systema naturae per regna tria naturae :secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (L. Salvius, Ed.; pp. 1–881). Impensis Direct. Laurentii Salvii. https://doi.org/10.5962/bhl.title.542

Lockyer, C. (1993). Seasonal Changes in Body Fat Condition of Northeast Atlantic Pilo Whales, and their Biological Signficance. Report of the International Whaling Commission (Special Issue), 14, 325–350.

Luque, S. P., & Ferguson, S. H. (2010). Age structure, growth, mortality, and density of belugas (Delphinapterus leucas) in the Canadian Arctic: Responses to environment? Polar Biology, 33(2), 163–178. https://doi.org/10.1007/s00300-009-0694-2

Mansfield, A. W., Smith, T. G., & Beck, B. (1975). The Narwhal, Monodon monoceros , in Eastern Canadian Waters. Journal of the Fisheries Research Board of Canada, 32(7), 1041–1046. https://doi.org/10.1139/f75-122

Merella, M., Collareta, A., Granata, V., Casati, S., & Bianucci, G. (2022). New remains of Casatia thermophila (Cetacea, Monodontidae) from the Lower Pliocene arine vertebrate-bearing locality of arcille (Tuscany, Italy). Rivista Italiana Di Paleontologia e Stratigrafia, 128(1), Article 1. https://doi.org/10.54103/2039-4942/15459

Mikhalev, Y. A. (2019). Analysis of the Correlation Between Whale Length and Weight. In Whales of the Southern Ocean: Biology, Whaling and Perspectives of Population Recovery (pp. 31–62). Springer International Publishing. https://doi.org/10.1007/978-3-030-29252-2_2

Mitchell, E. D., & Reeves, R. R. (1981). Catch History and Cumulative Catch Estimates of Initial Population Size of Cetaceans in the Eastern Canadian Arctic. Report of the International Whaling Commission, 31, 645–682.

Ognetov, G. N. (2007). Половой диморфизм белухи Delphinapterus leucas [Sexual Dimorphism in the Beluga]. Problems of Fisheries, 8(2 (30)), 172–183.

Pallas, P. S. (1776). Reise durch verschiedene Provinzen des Rußischen Reichs (Vol. 3). Kayserliche Academie der Wissenschaften.

Pluskowski, A. (2004). Narwhals or Unicorns? Exotic Animals as Material Culture in Medieval Europe. European Journal of Archaeology, 7(3), 291–313. https://doi.org/10.1177/1461957104056505

Rice, D. W., & Wolman, A. A. (1971). The life history and ecology of the gray whale (Eschrichtius robustus). The American Society of Mammalogists. https://portals.iucn.org/library/node/17576

Sergeant, D. E. (1973). Biology of White Whales ( Delphinapterus leucas ) in Western Hudson Bay. Journal of the Fisheries Research Board of Canada, 30(8), 1065–1090. https://doi.org/10.1139/f73-178

Sergeant, D. E., & Brodie, P. F. (1969). Body Size in White Whales, Delphinapterus leucas. Journal of the Fisheries Research Board of Canada, 26(10), 2561–2580. https://doi.org/10.1139/f69-251

Shpak, O., Meschersky, I., Glazov, D., Litovka, D., Kuznetsova, D., & Rozhnov, V. (2020). Structure and Assessment of Beluga Whale, Delphinapterus leucas, Populations in the Russian Far East. Marine Fisheries Review, 81, 72–86. https://doi.org/10.7755/MFR.81.3-4.3

Skovrind, M., Castruita, J. A. S., Haile, J., Treadaway, E. C., Gopalakrishnan, S., Westbury, M. V., Heide-Jørgensen, M. P., Szpak, P., & Lorenzen, E. D. (2019). Hybridization between two high Arctic cetaceans confirmed by genomic analysis. Scientific Reports, 9, 7729. https://doi.org/10.1038/s41598-019-44038-0

Suydam, R. S. (2009). Age, growth, reproduction, and movements of beluga whales (Delphinapterus leucas) from the eastern Chukchi Sea, Robert Scott Suydam [Doctoral dissertation]. University of Washington School of Aquatic and Fishery Sciences.

Tomilin, A. G. (1967). Mammals of the USSR and Adjacent Countries Volume IX: Cetacea (Kitoobraznye). Israel Program of Scientific Translations.

Tyrrell, M. (2008). Nunavik Inuit Perspectives on Beluga Whale Management in the Canadian Arctic. Human Organization, 67(3), 322–334.

Vélez-Juarbe, J., & Pyenson, N. D. (2012). Bohaskaia monodontoides , a new monodontid (Cetacea, Odontoceti, Delphinoidea) from the Pliocene of the western North Atlantic Ocean. Journal of Vertebrate Paleontology, 32(2), 476–484. https://doi.org/10.1080/02724634.2012.641705

Víkingsson, G., Sigurjónsson, J., & Gunnlaugsson, T. (1988). On the Relationship Between Weight, Length and Girth Dimensions in Fin and Sei Whales Caught off Iceland. Rep. Int. Whal. Commn, 38, 323–326.

Vladykov, V. D. (1944). Études sur les mammifères aquatiques. III. Chasse, biologie et valeur économique du marsouin blanc ou béluga (Delphinapterus leucas) du fleuve et du golfe du Saint-Laurent (p. 194). Département des pêcheries. Province de Québec.

Worm, O. (1655). Museum Wormianum; seu, Historia rerum rariorum, tam naturalium, quam artificialium, tam domesticarum, quam exoticarum, quae Hafniae Danorum in a aedibus authoris servantur. ex officina Elseviriorum. https://doi.org/10.5962/bhl.title.119357

Author: CallmeJoe

College Graduate in Biology who's primarily a fan of Godzilla and other properties.

Leave a comment